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Full text of "Bulletin of the Natural History Museum Entomology"

ISSN 0968-0454 



Bulletin of 
The Natural History Museum 



Entomology Series 




VOLUME 63 NUMBER 1 23 JUNE 1994 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British 
Museum (Natural History) ), instituted in 1949, is issued in four scientific series, 
Botany, Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Entomology Series is produced under the editorship of the 

Keeper of Entomology: Dr R.P. Lane 

Editor of Bulletin: Dr P.C. Barnard 



Papers in the Bulletin are primarily the results of research carried out on the 
unique and ever-growing collections of the Museum, both by the scientific staff 
and by specialists from elsewhere who make use of the Museum's resources. 
Many of the papers are works of reference that will remain indispensable for 
years to come. All papers submitted for publication are subjected to external 
peer review before acceptance. 

A volume contains about 192 pages, made up by two numbers, published in the 
Spring and Autumn. Subscriptions may be placed for one or more of the series 
on an annual basis. Individual numbers and back numbers can be purchased and 
a Bulletin catalogue, by series, is available. Orders and enquiries should be sent 
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Intercept Ltd. 
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Wo rid List abbreviation: Bull. nat. Hist. Mus. Lond. (Ent.) 
© The Natural History Museum, 1994 



Entomology Series 
ISSN 0968-0454 Vol. 63, No. 1, pp. 1-136 

The Natural History Museum 

Cromwell Road 

London SW7 5BD Issued 23 June 1994 

Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press, Oxford 



Bull. nai. Hist. Mus. Lond. (Ent.) 63(1):1-136 Issued 23 June 1994 

A revision of the Indo-Pacific 
species of Ooencyrtus 
(Hymenoptera: Encyrtidae), 
parasitoids of the immature stages 
of economically important insect 
species (mainly Hemiptera and 
Lepidoptera) 

D.-W. HUANG & J.S. NOYES* 

Institute of Zoology, Chinese Academy of Sciences, Beijing, P.R. China 

* Department of Entomology, The Natural History Museum, Cromwell Road, 

London SW7 5 BD 

CONTENTS 



Synopsis 1 

Introduction 2 

Biology of Ooencyrtus spp 2 

Synopsis of hosts of Indo-Pacific species 4 

Use of Ooencyrtus in biological control 7 

Depositories 7 

Acknowledgements 7 

Systematic Section 8 

Ooencyrtus Ashmead 8 

Comments on generic synonymy 9 

Generic diagnosis 9 

Systematic relationships within the genus 10 

Identification of species 10 

Abbreviations used in text 11 

Key to Oriental species of Ooencyrtus (females) 1 1 

Review of species 16 

Excluded species 83 

References 84 

Illustrations 88 

Index to scientific names 134 



Synopsis. This revision treats 70 species of the genus Ooencyrtus found in the 
Indo-Pacific region, including south China, but excluding Australia and New Zealand. 
One new generic and 7 new specific synonymies are proposed, 49 species are described 
as new 'and 11 lectotypes are designated. Each species is characterized by a diagnosis 
and/or full species description, its known distribution and hosts are reviewed, and a key 
is provided to the females. 

Correspondence to Dr J.S. Noyes 



©The Natural History Museum. 1994 



D. -W.HUANG AND J.S. NOYES 



INTRODUCTION 



The genus Ooencyrtus is of particular interest 
because it is widespread and species-rich in all 
geographic regions and its species parasitise 
diverse hosts and host stages. Many species para- 
sitise immature stages of pest species and thus 
have potential value as biological control agents. 
Some species also act as minor pests by parasitis- 
ing eggs of butterflies being reared for commer- 
cial purposes in Malaysia and Thailand. 

This interest is reflected by the relatively large 
amount of material of the genus that is received 
by the identification services of the The Natural 
History Museum and International Institute of 
Entomology from the Indo-Pacific region. 
Unfortunately, identification is difficult because 
the taxonomy of the genus in this part of the 
world is inadequate. This revision was under- 
taken as a first step towards improving this 
situation and it is perhaps not surprising that of 
the 70 species treated here, more than two-thirds 
are described as new. 



BIOLOGY OF OOENCYRTUS 



the solitary eggs of Sphingidae (Battisti et ai, 
1988). At least one species is known to be a 
gregarious parasitoid of the solitary eggs of but- 
terflies, e.g. papilionis (see p. 79). Two species 
are known to attack different stages of hosts in 
different orders. Ooencyrtus kuvanae, has been 
recorded both as a primary solitary parasitoid of 
the eggs of gypsy moth (Lymantria dispar) and as 
a gregarious hyperparasitoid of the prepupal and 
pupal stages of the moth's braconid primary 
parasitoid. Muesebeck & Dohanian (1927) found 
that female kuvanae from field-collected gypsy 
moth eggs would oviposit into the immature 
stages of Apanteles melanoscelus within their 
cocoons after they had emerged from the gypsy 
moth caterpillar. Up to 17 individuals subse- 
quently issuing from a single braconid host. The 
same species of Ooencyrtus has also been 
recorded as a primary parasitoid of Anastatus 
bifasciatus Fonscolombe in gypsy moth eggs 
(Howard & Fiske, 1911), although Crossman 
(1925) stated that he knew of only one such case 
in 14 years of working with these species. Ooen- 
cyrtus submetallicus (Howard) is best known as a 
primary parasitoid of the heteropterous eggs, but 
it has also been recorded as a parasitoid of the 
pupae of a chloropid dipteran (Legner & Bay, 
1965a,b). 



Host range 

The majority of species are parasitoids of the 
eggs of Hemiptera or Lepidoptera, but species 
are known that attack the prepupae of Lepi- 
doptera, braconid primary parasitoids of cater- 
pillars (Lepidoptera), immature stages of 
Dryinidae attacking auchenorrhynchous 
Homoptera, nymphal stages of Aphididae 
(Homoptera), immature stages of Syrphidae 
(Diptera) or Coccinellidae (Coleoptera) feeding 
on aphids, or the pupae of Chloropidae 
(Diptera). 

Host specificity 

The degree of host specificity is not known with 
any certainty. Some species appear to act only as 
solitary parasitoids of lepidopterous eggs laid in 
batches, such as pinicolus which parasitises the 
eggs of Lymantriidae and Lasiocampidae (see 
Trjapitzin, 1989). Other species may be parasi- 
toids of the eggs of Heteroptera or Lepidoptera 
which are laid in batches, e.g. pityocampae which 
will parasitize the eggs of Lasiocampidae, Not- 
odontidae (Lepidoptera) as well as Coreidae and 
Pentatomidae (Hemiptera), although the same 
species has also been recorded as a parasitoid of 



Ovary development 

Adult females emerge from the host with unde- 
veloped ovaries and the preoviposition period 
may vary from 1.5 to nearly four days depending 
on temperature (Tracy & Nechols, 1988). 

Mating 

Mating has been described in detail only for 
kuvanae (Brown, 1984). Females may mate sev- 
eral times, but if they participate in a post-mating 
ritual they will mate only once. The post-mating 
ritual involves the pair facing each other and the 
male touching the female with his antennae, fore 
legs and head. The female touches the male with 
her mouthparts at this time. Newly emerged, 
mating females may be swarmed over by males if 
populations are large. Mating males appear to 
clean the female with their mouthparts, they may 
also try to mate with a female before her wings 
are fully expanded. 

Host finding and oviposition 

Evidence suggests that, in at least two species 
[fecundus and pityocampae), female adult parasi- 
toids are attracted to chemicals or odours pro- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



duced by the adult female host (Laraichi & 
Voegele, 1975; Battisti, 1989). This may be the 
result of the parasitoid being conditioned by 
encountering suitable freshly laid eggs after the 
mass emergence of a suitable host species (Bat- 
tisti, 1989). Oviposition may take some time 
depending upon the experience of the parasitoid. 
In kuvanae experienced females may take only 
five minutes to deposit an egg after the initial 
encounter whilst inexperienced females may take 
nearly twice as long (Lee & Lee, 1989), whilst in 
gregarious species oviposition may take nearly an 
hour per single host, e.g. johnsoni (Maple, 
1937). Host feeding, by the adult parasitoid, has 
been observed in at least two species (kuvanae, 
Lee & Lee, 1989; johnsoni. Maple, 1937) and is 
almost certainly common throughout the genus. 

Fecundity 

The species of host egg can influence fecundity 
(Laraichi, 1978a). Host density does not affect 
fecundity but lower host density increases the 
rate of superparasitism in solitary parasitoids 
(Laraichi, 1978b). The maximum overall fecun- 
dity may be as high as 200 (Crossman, 1925). In 
species that parasitise egg clusters the number of 
eggs deposited by the parasitoid will be limited to 
the number of mature eggs present in the ova- 
ries, in nezarae this averages about 17 (Takasu & 
Hirose, 1991). In kuvanae this normally results in 
a lower percentage parasitism of larger egg 
masses resulting from egg limitation (Weseloh, 
1972; Williams et al., 1990). Percentage parasit- 
ism in kuvanae may be as high as 80%, but is 
generally around 10-40%. The substrate on 
which the host eggs are laid may affect rate of 
parasitism. In kuvanae, the eggs of its host on red 
maple (smooth bark) have a higher rate of para- 
sitism than those on oak (rough bark) (Bellinger 
etal., 1988). 

Immature stages 

After oviposition the egg stalk remains protrud- 
ing through the chorion of the host egg. The 
larva remains attached to its egg shell for the first 
three instars and is metapneustic (Maple, 1937), 
being able to utilise atmospheric air directly 
through the protruding part of the egg (Maple, 
1937). The number of larval instars recorded 
varies from three (Gerling et al., 1976 in trin- 
idadensis; Matteson, 1981 in utetheisae; Cross- 
man, 1925 in kuvanae), four (Maple, 1937 in 
johnsoni; Laraichi, 1977 in fecundus, nigerrimus 
and telenomicida) to five (Parker, 1933 in kuva- 
nae; Takasu & Hirose, 1989 in nezarae). The true 



number of instars is probably four or five since 
these counts are based on the mandibles of the 
exuviae of previous instars which remain 
attached to the anal shield of the final instar 
larva. 

Development time 

In both kuvanae and anasae development from 
egg to adult takes about 18-35 days depending on 
temperature (Muesebeck & Dohanian, 1927; 
Tracy & Nechols, 1987). In manii no develop- 
ment of immatures occurs if the temperature is 
over 32.5°C (Rahim et al., 1991). O kuvanae has 
4 or 5 generations per year in more northerly 
parts of the USA (Crossman, 1925) and up to 7 
generations per year in Italy (Prota, 1966) and 
overwinters as an adult. 

Adult longevity 

In manii longevity of both sexes decreases with 
an increase in temperature, males will live 4 to 
1.2 days with temperatures of 15-36°C whilst 
females live from 10-1.7 days at the same tem- 
peratures. In general, longevity increases with an 
increase in humidity, with optimum at 50-70% 
RH (Rahim et al., 1991). This is also reflected by 
longevity and fecundity of the same species dur- 
ing different seasons, the highest fecundity (37 
eggs/female) being noted in April and the lowest 
(0.8 eggs/female) being noted in January and 
February (Yadav & Chaudhary, 1984). In some 
species day length may effect reproduction and 
longevity. For instance, in kuvanae longer day 
length results in more progeny but lower longev- 
ity (Weseloh, 1986). 

Sex ratio and sex determination 

The sex ratio varies from about to 1:1 to 4:1 in 
favour of females (johnsoni Maple, 1937). In 
anasae the proportion of females increases with 
number of hosts parasitized (Tracy & Nechols, 
1987). Sex ratio may also be influenced by tem- 
perature. For instance, in fecundus (Laraichi, 
1978c) all progeny are female if the ovipositing 
female is subjected to temperatures of 30°C but 
all progeny are male if the ovipositing female is 
subjected to a temperature of 35°C (Laraichi, 
1978c). In O. submetallicus similar temperatures 
produce similar results with hermaphrodite 
mosaics being produced if the developing prog- 
eny are subjected to intermediate temperatures 
(Wilson, 1962; Wilson & Woolcock, 1960). This 
suggests that the sex of the offspring may be 
under the control of a microorganism similar to 



D. -W.HUANG AND J.S. NOYES 



that found in some species of Trichogramma (see 
Stouthamer et al, 1990; Stouthamer, 1990, 
1991). However, Kamay (in Brown, 1984) 
reported that if developing kuvanae were 
exposed to temperatures of around 35°C this 



resulted in a higher proportion of females, possi- 
bly because of higher male mortality. Seasonal 
fluctuations of temperature influence the sex 
ratio oimanii (Yadav & Chaudhary, 1984). 



SYNOPSIS OF HOSTS OF INDO-PACIFIC SPECIES 



[] extralimital records 

* probably incorrect host association 

! laboratory reared 

(H) hyperparasitoid 

All reared from eggs except those prefixed by (L) - from larvae, and (P) - from prepupae or pupae. 



HOST 



PARASITOID 

(Ooencyrtus sp.) 



Unknown eggs 

COLEOPTERA 

Chrysomelidae 

Plesispa reichei Chapius 

Podontia quatuordecimpunctata (Linnaeus) 

Podontia quatuordecimpunctata (Linnaeus) 

Coccinellidae 
(P)indet. 

DIPTERA 

Syrphidae 

(P)Allograpta exotica (Wiedemann) 
(L)Dideopsis pura (Curran) 
(L)[Ischiodon scutellaris (Fabricius) 
(L)[Ischiodon aegyptius Wied (Wiedemann) 
(L)Paragus auritus Stuckenberg 

HEMIPTERA 

indet family 

Heteroptera 

Alydidae 
Leptocorisa sp. 
Leptocorisa acuta Thunberg 
[Mirperus jaculus (Thunberg) 
Piezodorus hybneri (Fabricius) 
Rip tortus sp. 
Rip tortus sp. 
[Riptortus dentipes (Fabricius) 

Coreidae 
Amblypelta sp. 
Amblypelta cocophaga China 
Amblypelta lutescens Distant 
Amblypelta papuensis Brown 

[Anoplocnemis curvipes (Fabricius) 



pindarus 

corbetti 

podontiae 

guamensis 



guamensis 
guamensis 
guamensis] 
guamensis] 
guamensis 



ferrierei 



utetheisae 

utetheisae 

utetheisae] 

utetheisae 

cybele 

utetheisae 

utetheisae] 



caurus 

ilion 

caurus 

utetheisae 

caurus 

utetheisae] 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



[Clavigralla elongata Signoret 
[Clavigralla tomentosicollis Stal 
Dasynus kalshoveni Blote 
Dasynus piperis China 
[Gonocerus acutangulatus Goeze 
[Gonocerus juniperi Herich-Schaeffer 
Mictis profana (Fabricius) 
[Pseudotheraptus wayi Brown 

Pentatomidae 

indet. 

[Brachynema germarii (Kolenati) 

[Dolycoris penicillatus Horvath 

Eocanthecona furcellata (Wolff) 

Nezara viridula (Linnaeus) 

Plataspidae 

Brachyplatys pacificus (Dallas) 

Cratoplatys sp. 

Scutelleridae 

[Eurygaster integriceps Puton 

[Aelia sp. 

Tessaratomidae 
Tessaratoma papillosa (Drury) 
Tessaratoma javanica (Thunberg) 
Pycanum ponderosum Stal 

Homoptera 
indet. family 

Lophopidae 
Pyrilla spp. 
Pyrilla perpusilla Walker 

Aphididae 
(L)indet. 



utetheisae] 

utetheisae] 

icarus 

utetheisae 

\telenomicida] 

telenomicida] 

utetheisae 

utetheisae] 



lucens 

telenomicida] 

telenomicida] 

iulus 

utetheisae 

pacificus 
ceres 



telenomicida] 
telenomicida] 



phongi 
phongi 
phongi 

midas 

manii 
manii 

pallidipes * 



LEPIDOPTERA 

indet. 
indet. 
indet. 
(P)indet. leafmining family 

Agonoxenidae 

(L)Agonoxena pyrogramma Meyrick 

Arctiidae 

[Utetheisa pulchella (Linnaeus) 

Bombycidae 

Rondotia menciana Moore 

Crambidae 

Chilo terenellus Pagenstecher 

Danaidae 

Danaus chrysippus (Linnaeus) 
Euploea core (Cramer) 
Tirumala limniace (Cramer) 

Epipyropidae 

(P)Epiricania melanoleuca (Fletcher) 

Gracillariidae 

(P)Acrocercops globulifera Meyrick 

(P)Conopomorpha cramerella (Snellen) 



phongi * 
javanicus 
utetheisae 
ooii 

shakespearei 

utetheisae 

hercle 

papilionis 

papilionis 
papilionis 
papilionis 



oott 
ooii 



D. -W.HUANG AND J.S. NOYES 



Heliconiidae 

Heliconius charitonius (Linnaeus) 

Hesperiidae 

Erionota thrax (Linnaeus) 

Hasora sp. 

Lasiocampidae 

Dendrolimus sp. 

Dendrolimus kikuchii Matsumura 

Dendrolimus spectabilis Butler 

Malacosoma americana 

Malacosoma neustria tartacea Motschulsky 

Taragama repanda Hubner 

Lymantriidae 

Aroa cometaris Butler 

Euproctis chrysorrhoea (Linnaeus) 

Hemerocampa leucostigma Abbot & Smith 

Hemerocampa definata Packard 

[Lymantria dispar (Linnaeus) 

[Lymantria fumida Butler 

Lymantria xylina Swinhoe 

[Nygmia phaeorrhoea Donovan 

[Stilpnotia salicis Linnaeus 

Noctuidae 

Achaea Janata (Linnaeus) 
(L)Exelastis atomosa (Walsingham) 
Othreis fullonia (Clerck) 
Othreis fullonia (Clerck) 
(L)Mythimna unipuncta (Haworth) 

Notodontidae 
Clostera cupreata (Butler) 
Stauropus lichenina Butler 
Turnaca acuta (Walker) 

Nymphalidae 

Ariadne ariadne (Linnaeus) 

Caligo memnon (Felder) 

Hypolimnas bolina Linnaeus 

Kallima sp. 

Phalanta phalantha (Drury) 

Junonia lemonias (Linnaeus) 

Tanaecia julii Bougainville 

Papilionidae 
Troides Helena Linnaeus 
Papilio sp. 
Papilio sp. 

Papilio aegeus Donovan 
Papilio agamemnon (Linnaeus) 
Papilio citri (?lapsus for demoleus 
Papilio demoleus Linnaeus 
Papilio helenus Linnaeus 
Papilio memnon Linnaeus 
Papilio polytes Linnaeus 
Papilio rumanzovia Eschscholtz 

Pieridae 

(L)Delias sp. 

Eurema sp. 

Hebomoia glaucippe (Linnaeus) 



papilionis 

pallidipes 
papilionis 

endymion 

endymion 

kuvanae 

\kuvanae 

kuvanae 

Uelenomicida 



papilionis 
\kuvanae 
\kuvanae 
Ikuvanae 
kuvanae] 
kuvanae] 
Ikuvanae 
kuvanae] 
kuvanae] 

lucens 

guamensis 

crassulus 

papilionis 

guamensis 

lucina 
bo re as 
macula 

papilionis 
pallidipes 
papilionis 
papilionis 
papilionis 
papilionis 
papilionis 

papilionis 

papilionis 

plautus 

papilionis 

papilionis 

plautus 

papilionis 

papilionis 

paplionis 

papilionis 

papilionis 

larvarum 
larvarum 
hera 



Saturniidae 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 

Attacus atlas Fabricius 
Callosamia promethea Drury 
Cricula sp. 

Cricula elaezia Jordan 
[Eriogyna pyretorum Westwood 
Hemileuca maia Drury 
Hemileuca oliviae Churchill 

Sphingidae 

indet. 

Amorpha populi austanti Staudinger 

Cephonodes hylas (Linnaeus) 

IGnathothlibas erotus eras (Boisduval) 

Hippotion celerio (Linnaeus) 

NEUROPTERA 

Myrmeleontidae 

indet. 



phoebi 

\kuvanae 

javanicus 

dione 

kuvanae] 

Ikuvanae 

Ikuvanae 



endymion 

Uelenomicida 

papilionis 

sphingidarum 

crassulus 



USE OF OOENCYRTUS IN 
BIOLOGICAL CONTROL 

Although species of Ooencyrtus undoubtedly 
play an important role in the regulation of the 
populations of many insect species worldwide, 
they have not proven to be of great value in pest 
control. Their use in classical biological control 
programmes is summarised in Table 1. Perhaps 
the best documented example is that of Ooencyr- 
tus kuvanae for the control of the gypsy moth 
(Lymantria dispar) in North America and 
Europe. Brown (1984) comments that although 
affording some control, the species is unlikely to 
be of much benefit by itself, although it probably 
causes a post outbreak collapse which in turn 
increases the length of the interval between out- 
breaks. 

A further species, Ooencyrtus fecundus Fer- 
riere & Voegele, may be of some benefit if used 
by means of inundative releases against het- 
eropterous pests of wheat in North Africa (Larai- 
chi & Voegele, 1975). 



DEPOSITORIES 

BMNH The Natural History Museum, London, 

England 
BPBM Bernice P. Bishop Museum, Hawaii 
CNC Canadian National Collection, Ottawa, 

Canada 
IARI Indian Agriculture Research Institute, New 

Delhi, India 
IEE Institito di Entomologfa Espanol, Madrid, 

Spain 
IRSN Institute Royal des Sciences Naturelles de 

Belgique, Brussels, Belgium 



IZAS Instiute of Zoology, Chinese Academy of 

Sciences, Beijing, PR. China 

MARI Malaysian Agricultural Research and 
Development Institute, Kuala Lumpur, 
Malaysia 

MNHN Museum National d'Histoire Naturelle, 
Paris, France 

MZB Museum Zoologicum Bogoriense, Bogor, 

Indonesia 

ORSTOM Office de la Recherche Scientifique et 
Technique Outre-Mer, Paris, France 

PPRI Plant Protection Research Institute, Preto- 

ria, South Africa 

OMB Queensland Museum, Brisbane, Australia 

TAMU Texas A&M University, Texas, USA 

USNM United States National Museum, Washing- 
ton, D.C., USA 

ZAMU Zoology Department, Aligarh Muslim Uni- 
versity, Aligarh, India 

ZISP Zoological Institute, St Petersberg, Russia 



ACKNOWLEDGEMENTS 

We thank The Royal Society which provided the 
senior author the opportunity to study at The 
Natural History Museum in London. Many 
thanks to Mr David Foster of The Royal Society, 
who gave the senior author consistent help dur- 
ing his stay in UK. The Natural History Museum 
made its collections, facilities and libraries avail- 
able to us for the project. The senior author's 
host institution, Institute of Zoology, Chinese 
Academy of Sciences permitted him one year off. 
We also thank Nigel Wyatt, Jeremy Holloway, 
Gary Stonedahl, Phil Ackery, Mike Shaffer and 
Mick Webb for checking host names, and thanks 
especially to John LaSalle for his valuable com- 
ments concerning the manuscript. Finally we are 
grateful to Dr M. Schauff (USNM), Dr G. 



8 D. -W.HUANG AND J.S. NOYES 

Table 1 A summary of the use of Ooencyrtus spp. in classical biological control programmes worldwide 
(Abbreviations: NE - not established; NR - not released; ? - no subsequent information; SC - successful 
control; P - partial control; E - established but no further information available; NC - established but no 
significant control achieved). 



Target pest species 



Introduced Ooencyrtus sp.; country; year; result; source 



HEMIPTERA 

Amblypelta cocophaga 

Amblypelta theobromae 
Anasa tristis 
Eurygaster integriceps 

Murgantia histrionica 
Nezara viridula 



Pseudotheraptus wayi 

Soybean stink bugs 
LEP1DOPTERA 
Ascotis selenaria 
Calpodes ethlius 

Conopomorpha cramerella 
Erionota thrax 

Hemerocampa leucostigma 
Hemileuca oliviae 
Lymantria dispar 



Othreis fullonia 



- sp.; Solomon Is; 1937-38; NE; Phillips (1941) 

- malayensis (?= utetheisae, misident.); Solomon Is; 1937-38; NE; Phillips (1941) 

- malayensis (?= utetheisae, misident.); Papua New Guinea; 71974; ?; Young 
(1982) 

- malayensis (?= utetheisae, misident.); USA (Massachusetts); 1981; ?; Coulson et 
al. (1988) 

-fecundus; USSR; ?; NR; Izhevskiy (1988) 

- nigerrimus; USSR; ?; NR; Izhevskiy (1988) 

- telenomicida; USSR; ?; NR; Izhevskiy (1988) 
-johnsoni; Bermuda; 1953; NE; Bennett & Hughes (1959) 
-johnsoni; Hawaii; 1940; E; Clausen in Clausen (1978) 
-johnsoni; Australia; 1953; NR; Wilson (1960) 

- malayensis (?= utetheisae, misident.); USA (Massachusetts); 1981; ?; Coulson et 
al. (1988) 

-submetallicus; Australia; 1952-57; NE; Wilson (1960) 

- submetallicus; Hawaii; 1962; NE; Davis (1964), Waterhouse & Norris (1987) 
-submetallicus; USA (Florida); 1973; ?; CIBC (1974) 

- trinidadensis; Hawaii; 1962; NE; Davis (1964), Waterhouse & Norris (1987) 
-sp. (?= utetheisae); Zanzibar; 1959; ?; Greathead (1971) 

-sp. (?= utetheisae); Kenya; 1959; ?; Greathead (1971) 
-nezarae; Brazil; 1983-1985; ?; Kobayashi & Cosenza (1987) 

- ennomophagus; Israel; 1977; NR; Wysoki (1979) 

- sp. (= calpodicus); Bermuda; 1953, 1962-63; NC; Cock (1985) 

- sp. (= calpodicus); St Vincent; 1950-51; E; Cock (1985) 

-sp. (= ooii); Malaysia (Sabah); 1987; NE; CIBC (1988, 1989, 1990) 
-pallidipes; Hawaii; 1973; SC; Mau, et al. (1980) 
-pallidipes; Mauritius: 1971-72; P; Waterhouse & Norris (1989) 
-kuvanae; USA; 1917, 1921; NE; Dowden (1962) 

- kuvanae; USA (New Mexico); 1913-16; NE; Clausen (1956) 

- kuvanae; Algeria; 1925-26, 1931; E; Lepigre (1932), Clausen (1978) 

- kuvanae; Canada (Ontario); 1976; E; Brown (1984) 

- kuvanae; Czechoslovakia; 1922; E; Clausen (1978) 

- kuvanae; Morocco; 1924-26; E; Brown (1984), Clausen (1978) 
-kuvanae; Portugal; 1932; E; Brown (1984), Clausen (1978) 
-kuvanae; Spain; 1923-27; E; Brown (1984), Clausen (1978) 
-kuvanae; USA; 1908-28, 1967-1971, 1981; E; Brown (1984), Peck (1963), 
Clausen (1956, 1978), Coulson et al. (1988) 

- kuvanae; CIS; 1987; ?; Volkov & Mirohova (1990) 

- kuvanae; Yugoslavia; 71960; ?; Brown (1984) 

- sp. (?= papilionis); American Samoa; ?; ?; Waterhouse & Norris (1987) 

- sp. (?= papilionis); Western Samoa; ?; ?; Waterhouse & Norris (1987) 



Nishida (BPBM), Dr J.-L. Nieves Aldrey (IEE), 
Dr I. Trjapitzin (ZISP), Dr G.L. Prinsloo 
(PPRI), Dr G. Gibson (CNC), Dr P. Dessart 
(IRSN) and Mr E.C. Dahms (QMB) for the loan 
or gift of material. 



SYSTEMATIC SECTION 



OOENCYRTUS Ashmead 

Ooencyrtus Ashmead, 1900: 381. Type species: 
Encyrtus clisiocampae Ashmead, by original 
designation. 

Echthrodryinus Perkins, 1906: 252. Type species: 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



Echthrodryinus destructor Perkins, by mono- 
typy. Synonymy with Ooencyrtus by Noyes & 
Hayat, 1984. 

Ectopiognatha Perkins, 1906: 254. Type species: 
Ectopiognatha minor Perkins, by designation 
of Gahan & Fagan, 1923: 49. Syn.nov. 

Schedius Howard, 1910: 2. Type species: Sche- 
dius kuvanae Howard, by original designation. 
Synonymy with Ooencyrtus by Ferriere, 1931. 

Tetracnemella Girault, 1915: 170. Type species: 
Tetracnemella australiensis Girault, by original 
designation. Synonymy with Ooencyrtus by 
Noyes & Hayat, 1984. 

Xesmatia Timberlake, 1920: 424. Type species: 
Xesmatia flavipes Timberlake, by original des- 
ignation. Synonymy with Ooencyrtus by Noyes 
& Hayat, 1984. 

Pseudolitomastix Risbec, 1954: 1068. Type spe- 
cies: Litomastix creona Risbec. Synonymy with 
Ooencyrtus by Annecke & Mynhardt, 1973. 

Comments on generic synonymy 

Species previously combined with the genus 
Ectopiognatha can be separated from Ooencyrtus 
by the quadridentate mandibles and a broadened 
and flattened scape. However, one of the species 
included in this revision (leander sp.n.) is 
extremely close in general structure to both spe- 
cies included in Ectopiognatha except that it has 
a cylindrical scape. In addition, several described 
species of Ooencyrtus have a scape which is 
distinctly broadened. The biology of Ectopiog- 
natha spp. (parasitoids of the eggs of Hemiptera) 
falls within the range of that found in Ooencyrtus 
(see below) and, therefore, we have no hesita- 
tion in treating the two genera as synonymous. 

Noyes and Hayat (1984) treated Echthrodryi- 
nus as a synonym of Ooencyrtus whilst Gordh & 
Trjapitzin (1978) and later Trjapitzin (1989) have 
treated it as valid. It is not possible to separate 
the two genera on morphological grounds and 
the only possible basis for continuing to treat 
them as distinct is that the included species have 
different biologies. Ooencyrtus could be 
restricted to primary parasitoids (or perhaps 
hyperparasitoids) of the eggs of various insects, 
whilst Echthrodryinus could include primary 
parasitoids or hyperparasitoids of the larvae or 
prepupal stages of certain holometabolous 
insects, e.g. Dryinidae, Braconidae and Gracilla- 
riidae. However, one species, Ooencyrtus kuva- 
nae, has been recorded both as a primary 
parasitoid of the eggs of gypsy moth (Lymantria 
dispar) and as a hyperparasitoid of the prepupal 
and pupal stages of the braconid Apanteles mel- 
anoscelus in their cocoons after they emerge 



from their gypsy moth caterpillar host (see 
below). Recent work (Noyes, 1985 and Prinsloo, 
1987) has shown that the host range of Ooencyr- 
tus spp. is much more diverse than had previ- 
ously been thought, many species being noted as 
parasitic on nymphs of Aphididae (Hemiptera) 
and Syrphidae (Diptera), even to the extent 
where specimens that attack dipterous or 
coleopterous larvae are morphologically indistin- 
guishable from those that attack heteropterous 
eggs (see Noyes, 1985). In view of this, we are 
continuing to treat Echthrodryinus as synony- 
mous with Ooencyrtus. 

Generic diagnosis 

In the most comprehensive classification of the 
Encyrtidae currently available, Trjapitzin (1973, 
1989) includes Ooencyrtus in the Encyrtinae, 
tribe Microteryini, subtribe Ooencyrtina. Unfor- 
tunately, this classification does not provide any 
meaningful diagnoses for the tribes or subtribes. 
We are therefore hopeful that the following 
diagnosis will enable species to be assigned cor- 
rectly to Ooencyrtus: 

Robust, squat species, never conspicuously 
slender and elongate; thorax with posterior 
margin of mesoscutum weakly to strongly con- 
vex medially and overlying axillae centrally so 
that when thorax is in normal resting position 
axillae appear to be widely separated; 
mesopleuron posteriorly expanded so that it 
touches base of gaster, or nearly so, and com- 
pletely conceals the metapleuron and propo- 
deum from lateral view immediately above the 
hind coxae; forewing with marginal vein punc- 
tiform or not much longer than broad, 

Species of Ooencyrtus can be most easily con- 
fused with Trichomasthus, Helegonatopus and 
Psyllaephagus. Both Helegonatopus and Psyl- 
laephagus have the mesopleuron normal, not 
expanded posteriorly and not touching the base 
of the gaster so that in lateral view the metapleu- 
ron and propodeum are not obscured and touch 
the hind coxae. Species of Trichomasthus are 
generally much larger, usually being about 1.5 
mm long, the marginal vein of the forewing is 
normally several times longer than broad, and all 
species are parasitoids of scale insects (Coc- 
coidea). 

Superficially, the placement of the genus 
within the tribe Microteryini may appear some- 
what questionable, especially on biological 
grounds. Morphologically, it is possibly to see an 
evolutionary trend from a Microterys-Mke or 
Trichomasthus-Wke ancestor, some species of 
Microterys and Trichomasthus being quite similar 



10 



D. -W.HUANG AND J.S. NOYES 



structurally and, as stated above, it can be some- 
times quite difficult to separate Ooencyrtus from 
Trichomasthus . If it is hypothesized that the 
plesiomorphic host association for the tribe is 
scale insects (Homoptera; Coccoidea) (see 
Trjapitzin, 1989) then it would be possible to 
envisage a switch from parasitising insects with 
hard a scale covering and a host immune system, 
to parasitising the eggs of other insects which 
have a hard outer shell and no immune system. A 
further switch to hyperparasitism of Braconidae 
or Dryinidae, or parasitism of the immature 
stages of holometabolous insects (Diptera and 
Lepidoptera) is much more difficult to envisage, 
but at least two species are known to parasitise 
both hemipterous or lepidopterous eggs and the 
prepupae or pupae of holometabolous insects, 
i.e. kuvanae (Muesebeck & Dohanian, 1927) and 
submetallicus (see Noyes, 1985). 

Systematic relationships within the 
genus 

The relationships within Ooencyrtus are unclear 
and we do not attempt to provide any sort of 
formal classificatory framework within the genus 
by proposing subgeneric categories. However, 
probable assemblages of closely related species 
within the genus are highlighted in the comments 
sections of the appropriate species. 

Identification of species 

The following works should be consulted as aids 
to the identification of species from outside the 
Oriental region: Peck (1963) and Gordh (1979) 
for North America; Noyes (1985) and De Santis 
(1988) for South America; Trjapitzin (1989) for 
the Palaearctic; Prinsloo (1987) for Africa. 

Previous work dealing with the species within 
the Indo-Pacific area has not much value, either 
because of inadequate coverage of the species, or 
because it relies on poor characters whilst over- 
looking important diagnostic features. The only 
key available (Trjapitzin et ah, 1978) is based 
almost entirely on inadequate original descrip- 
tions and is therefore of little use. 

Although we describe males of each species 
when they are available, they are excluded from 
the key to species. Males are often difficult to 
identify because they do not present many fea- 
tures and are unknown for most of the species 
included here. However, the presence of males 
in a reared series may help to confirm the separa- 
tion of some closely related species, e.g. sphingi- 
darum and papilionis. 

During the course of this work we have found 



the following characters especially helpful in 
identifying species: 

Mandibles - several basic types, a) with one 
tooth and a broad straight, sometimes minutely 
denticulate truncation (Figs 10, 18, 139, 188, 204, 
333, etc.), b) with one tooth and a broad, slightly 
convex, minutely denticulate truncation (Figs 
303), c) one tooth and a distinctly emarginate 
truncation, or with two teeth and a truncation 
(Figs 75, 143, 150, etc.), d) tridentate (Fig. 69, 
91, 268, etc.), all teeth subequal, e) one or two 
teeth and an oblique minutely denticulate trunca- 
tion, (Fig. 240), f) three acute, unequal teeth 
(Fig. 176), or g) four teeth (Fig. 46). 
Clava - a) clava with apex rounded and sensory 
area at extreme apex only (Figs 42, 47, 51, etc.) 
b) clava with sutures transverse and parallel, 
apex obliquely truncate and sensory area 
enlarged (Figs 9, 104, 196, etc.), or c) clava with 
sutures oblique, apex obliquely truncate and 
sensory area enlarged (Figs 16, 22, 28, 193, etc). 
Eyes - a) conspicuously hairy, or b) almost 
naked. 

Ocelli - relative distance of ocelli from occipital 
margin in terms of their own diameters. 
Antennal toruli - relative distance below eyes or 
above mouth margin in terms of their own 
lengths. 

Interantennal prominence - coloration purple or 
metallic green or blue green, etc. 
Scutellum - relative depth and type of sculpture 
in relation to mesoscutum. 
Forewing - relative density and distribution of 
setae on both dorsal and ventral surfaces of basal 
cell and on ventral surface of costal cell; linea 
clava closed or open; relative length of marginal 
and postmarginal veins in relation to stigmal. 
Legs - the coloration of coxae, femora and tibia 
seems to be reliable although there may be a 
slight amount of variation in the intensity of any 
brown areas on the femora and tibia. The colora- 
tion of the fore and hind coxae may vary from 
yellow to largely brown in some species. 
Gaster - the relative length may be unreliable 
because it may depend on the preservation tech- 
niques used. In air dried specimens the gaster 
collapses and may thus be relatively shorter than 
in Critical Point Dried specimens where the 
gaster remains inflated (see Gordh & Hall, 
1976). The shape of the last tergite (apically 
rounded, truncate or medially invaginated) may 
be useful as well as the shape of the hypopygium 
(rectangular, triangular, with long or short ante- 
rior lateral projection, relative size of posterior 
incision, distribution and relative density of 
setae). 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



11 



Ovipositor - other than the usual characters such 
as relative length of the gonostyli or overall 
length of the ovipositor in relation to the mid 
tibia we have found that the shape of the proxi- 
mal part of the second valvifer can be extremely 
useful (e.g. compare upper parts of Figs 225 and 
226). The length of the ovipositor is here taken as 
the combined length of the second valvifer 
together with the third valvula (gonostylus). 



Abbreviations used in text 

F1,F2, etc 



second 



First funicle segments, 
funicle segment, etc. 

OOL - Ocular-ocellar line, or the shortest 

distance between each posterior 
ocellus and the adjacent eye margin. 

POL - Posterior ocellar line, or the short- 

est distance between the two poste- 
rior ocelli. 

Key to Oriental species of Ooencyrtus 
(females) 

1 At least one pair of coxae black or brown, 
concolorous with mesopleuron; gaster entirely 
black or brown without any yellow or orange 
areas 2 

— All coxae yellow (occasionally fore coxae 
slightly infuscate); gaster frequently yellow or 
orange in part, although often entirely black or 
brown 

33 

2(1) All coxae and femora conspicuously black or 
brown (rarely femora yellow with inconspicu- 
ous brown markings but if so then clava has a 
broad, slightly oblique apical truncation, the 
funicle segments are quadrate or transverse, 
the scape is subcylindrical (Fig. 9) and the 
scutellum has relatively shallow sculpture 
basally (Fig. 12)) 3 

— Not all coxae and femora black or brown, at 
least some of them yellow (clava usually with 
apex rounded but if with a distinct, slightly 
oblique apical truncation, then either some 
funicle segments are clearly longer than broad, 
or the scape is strongly flattened and only 
about 2.5 times as long as broad (Fig. 104), or 
the scutellum has deep punctate-sculpture 
basally (as in Fig. 109)) 20 

3(2) Antenna (Figs 1, 9, 16, 22, 28, 34) with clava 
robust and with a large sensory apical area 
which gives it a distinct transversely or 
obliquely truncate appearance 4 

Antenna (Figs 37, 42, 47, 51, 54, 59, 64, 67, 68, 
76, 83) with clava normally comparatively slen- 
der and with only a small apical sensory area 



which gives it a rounded or even pointed 
appearance 9 

4(3) Sensory part of clava less than one third length 
of clava (Figs 1,4,9) 5 

— Sensory part of clava more than one-third 
length of clava (Figs 16, 22, 28,34) 6 

5(4) Mesoscutum with numerous silvery setae; 
scutellum conspicuously reticulate throughout 
(Fig. 2); [fore-tibia of male characteristic (Fig. 

7)] 

guamensis (p. 16) 

Mesoscutum without silvery setae; posterior 
half of scutellum smooth (Fig. 12) ... acca (p. 
17) 

6(4) Antenna with first funicle segment at least 1.5 
times as long as broad and at least a little 
longer than sixth (Fig. 16) .... phongi (p. 18) 

— First funicle segment not, or hardly, longer than 
broad and shorter than sixth (Figs 22, 28, 34) . 7 

7(6) Frontovertex more than one-fifth head width; 
gaster apically acute with ovipositor exserted, 
the exserted part about as long as mid tibial 
spur; mandibles tridentate acus (p. 19) 

— Frontovertex less than one-fifth head width; 
gaster apically truncate with ovipositor more 
or less hidden; mandibles with one small tooth 
and a very broad truncation 8 

8(7) Scutellum with apical one-third smooth and 
very shiny caurus (p. 21) 

— Reticulate sculpture of scutellum extending 
almost to posterior margin with only a very 
narrow smooth and shiny marginal strip- 

lucens (p. 22) 

9(5) Forewing with linea calva closed posteriorly 
(Figs 38, 40); antenna with all funicle segments 
longer than broad (Fig. 37) 10 

— Either linea calva open posteriorly (Figs 44, 
48, 53, 55, 60, 65, 70, 77) or antenna with some 
segments quadrate or transverse (Figs 42, 83) 

11 

10(9) Flagellum bicolorous, funicle segments brown 
or testaceous, clava yellow .. mine rv a (p. 22) 

Flagellum unicolorous icarus (p. 23) 

11(9) Scutellum anteriorly with shallow, indistinct 
sculpture, the posterior part smooth; fron- 
tovertex at least one-third head width (Figs 43, 
45); mandibles (Fig. 46) with four teeth; ovi- 
positor exserted, the exserted part at least 
one-quarter as long as gaster 12 

— Scutellum with conspicuous, raised, regular 
reticulations, only margin narrowly and some- 
times apex smooth (Figs 71, 80); frontovertex 



12 



D. -W.HUANG AND J.S. NOYES 



normally distinctly less than one-third head 
width; mandibles tridentate (Figs 56, 74, 85) or 
with one or two teeth and a truncation (Figs 
69, 75); ovipositor not exserted, but if so then 
exserted part less than one-fifth length of 
gaster 13 

12(11) Antenna (Fig. 42) with Fl-4 subequal and 
clearly smaller than F5-6, clava as long as 
F3-6; exserted part of ovipositor half to one- 
third gaster length adonis (p. 24) 

— Antenna (Fig. 47) with Fl the smallest and 
much smaller than F2-6 which are subequal in 
size, clava about as long as F4-6; exserted part 
of ovipositor about one-quarter as long as 
gaster 
aeneas (p. 25) 

13(11) Visible part of ovipositor sheaths white, yellow 
or amber and extending past apex of last 
tergite, although occasionally hardly so ... 14 

— Visible part of ovipositor sheaths brown to 
dark brown, frequently not extending past 
apex of last tergite 15 

14(13) Reticulate sculpture of scutellum relatively 
shallow and hardly deeper than that on mesos- 
cutum, especially in apical half or so; mid 
femur and tibia hardly marked with brown, 
almost completely yellow; eye with conspicu- 
ous translucent setae phoebi (p. 26) 

— Reticulate sculpture of scutellum clearly 
deeper than that on mesoscutum except on 
sides an extreme apex; mid femur and tibia 
mostly dark brown; eye almost naked, with 
setae very short and inconspicuous 
kuvanae (p. 26) 

15(13) Scutellum uniformly bright green and with 
deep, more or less regular reticulate sculpture; 
mandibles with three acute teeth . dis (p. 27) 

— Scutellum green only at sides and apex, other- 
wise blue, purple or coppery and relatively dull 
with sculpture elongate-reticulate or striate 
towards sides; mandibles with one or two teeth 
and a truncation (Figs 69, 75) 16 

16(15) Frontovertex about one-third head width (Fig. 
62) hercle (p. 28) 

— Frontovertex not more than one-quarter head 
width (Fig. 74, 85) 17 

17(16) Clava relatively short, only about as long as 
F4-6 combined (Fig. 64) corbetti (p. 29) 

— Clava at least as long as F3-6 combined ... 18 

18(17) Mid and hind tibiae completely yellow, with- 
out a brown subbasal ring . podontiae (p. 30) 

Mid and hind tibiae each with a narrow brown 
subbasal ring 19 



19(18) Eye as broad as long or broader; antennae 
(Fig. 76) with Fl-2 subequal in size and 
smaller than F3-6 pindarus (p. 30) 

— Eye distinctly longer than broad; antennae 
(Fig. 83) with Fl-3 subequal in size and con- 
spicuously smaller than F4-6 . plautus (p. 31) 

20(2) Scutellum entirely smooth and shiny except for 
a small sculptured triangular area posterior to 
axillae, this extending less than half way along 
scutellum (Fig. 105); hypopygium reaching 
apex of gaster; gaster almost always with 
extensive yellow areas 21 

— Scutellum more extensively sculptured with 
sculpture discernible in apical half (Figs 106, 
109, 146); hypopygium never reaching apex, 
generally not reaching two-thirds along gaster; 
gaster without any yellow areas 23 

21(20) Forewing with a transverse fuscous band from 
marginal vein (Fig. 90); all coxae black- 
brown daphne (p. 33) 

— Forewing completely hyaline (Figs 98, 99); at 
least hind coxae yellow flavipes (p. 34) 

22(20) All coxae black or dark-brown 23 

— Fore-coxae yellow 30 

23(22) Scape strongly broadened and flattened, only 
about 2.5 times as long as broad; antennae 
strongly clavate with club extremely large (Fig. 
104) dione(p. 34) 

— Scape subcylindrical, at least about 4 times as 
long as broad; antenna not strongly clavate and 
with club of normal proportions, not conspicu- 
ously enlarged (Figs 108, 113, 120, 125, 126, 
129,134,141,142,148) 24 

24(23) Forewing with a median fuscous band (Fig. 107); 
antennae white except for radicle . dry as (p. 35) 

— Forewing hyaline; antennae testaceous or 
brown, not white 25 

25(24) Scutellum with relatively deep punctate- 
reticulate sculpture over more than half of its 
surface, this conspicuously deeper than sculp- 
ture of mesoscutum (Fig. 109) 26 

— Sculpture of scutellum not deeply punctate and 
generally not deeper than that on mesoscutum, 
but if so then deeper sculpture present only in 
basal one-third or less 27 

26(25) Clava paler than funicle, except perhaps F6; all 
funicle segments longer than broad, funicle 
clearly longer than scape and radicle together 
(Fig. 108) ceres (p. 36) 

— Clava concolorous with funicle or darker; at 
least four funicle segments quadrate or trans- 
verse, funicle shorter than scape and radicle 
together (Fig. 113) ilion (p. 37) 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



13 



27(26) Ovipositor sheaths yellow or pale orange, usu- 
ally protruding very slightly past apex of last 
tergite of gaster 28 

— Ovipositor sheaths dark brown, occasionally 
completely hidden by last tergite of gaster . 29 

28(28) Antennae relatively slender, the pedicel and 
flagellum together longer than head width and 
apical funicle segments longer than broad 
(similar to Fig. 108); fore tibiae yellow marked 
with pale brown iris (p. 38) 

— Antennae clavate, the pedicel and flagellum 
together shorter than head width and apical 
funicle segments transverse Or quadrate (Fig. 
120); fore tibiae almost entirely black or very 
dark brown javanicus (p. 39) 

29(28) Frontovertex not, or hardly, more than about 
one-quarter head width (Fig. 127); ventral 
surface of costal cell with only one complete 
row of stout setae (Fig. 128); dorsum of thorax 
moderately metallic with contrasting blue, 
green or purple areas iulus (p. 40) 

Frontovertex about one-third head width (Fig. 
132); ventral surface of costal cell with two or 
three complete rows of fine setae (Fig. 130); 
dorsum of thorax uniform dull green or green- 
ish blue without any strongly contrasting 
purple areas ixion (p. 40) 

30(22) Scutellum with at least posterior one-fourth 
smooth and shiny 31 

— Scutellum with less than posterior one-fifth 
smooth and shiny 32 

31(30) Anterior half of scutellum with regular raised 
reticulations, posterior half smooth and 
mirror-like; ocelli forming an angle of only 
slightly less than 90°; posterior femora mostly 
yellow but dorsally with a very narrow dark 
brown edge; ovipositor sheaths yellowish; 
mandibles with one obtuse small tooth and a 
broad truncation which bears five or six minute 
denticles (Fig. 137) 
elissa (p. 42) 

— Anterior three-quarters of scutellum elon- 
gately reticulate, posterior one-quarter 
smooth; ocelli very nearly forming an equilat- 
eral triangle; posterior femora mostly brown; 
ovipositor sheaths dark brown; mandibles 
without denticles on the truncate part (Fig. 
139) egeria {p. 43) 

32(30) All femora yellow, mid and hind coxae dark 
brown; scutellum reticulate except for a poste- 
rior mirror-like portion which is clearly delim- 
ited posteriorly by a sharply impressed line 
(Fig. 146) erebus (p. 44) 

— Hind femora at least partly brown, mid coxae 
more or less yellow; scutellum reticulate but 
without a clearly delimited mirror-like poste- 



rior portion endymion (p. 45) 

33(1) Gaster completely black, without any yellow 
or orange areas 34 

— Gaster partly yellow or orange, at least basally 
yellowish in ventral view 51 

34(33) Thorax with yellow or orange areas 35 

— Thorax completely dark, without any yellow or 
orange areas 36 

35(34) Clava slightly enlarged, clearly broader than 
funicle (Fig. 154) and truncated at apex with a 
conspicuous sensory area; mesoscutum with at 
least anterior margin orange; axillae orange; 
body larger (1.35 mm.) hymen (p. 47) 

— Clava not conspicuously enlarged and hardly 
broader than F6 (Fig. 160), its apex rounded 
and without a conspicuous sensory area; 
mesoscutum and axillae totally blue-green; 
body smaller (less than 1 mm.) . belus (p. 48) 

36(34) Scutellum with at least posterior one-fifth 
smooth and shiny 37 

Scutellum completely reticulate except for a 
very narrow marginal strip 41 

37(36) Flagellum bicolorous. clava yellow or pale 
yellow and contrasting with the brown to dark 
brown funicle segments 38 

Flagellum unicolorous, testaceous to dark 
brown 39 

38(37) Scape yellowish and clearly paler than funicle; 
F4-6 quadrate or transverse; clava as long as 
F4-6 together (Fig. 167); forewing hyaline 
bacchus (p. 49) 

Scape very dark brown and concolorous with 
funicle; F4-F6 longer than broad; clava as long 
as F3-F6 together (Fig. 169); forewing usually 
infuscate below marginal vein ... pacificus (p. 
50) 

39(37) Frontovertex about one-third head width; 
antenna with Fl transverse and much smaller 
than F2-5 which are also transverse (Fig. 175) 
larvarum (p. 50) 

— Frontovertex about one-fifth head width; 
antenna not with Fl conspicuously smaller 
than those that follow and at least some funicle 
segments longer than broad (Figs 181, 
187) 40 

40(39) Ovipositor sheaths yellow; forewing (Figs 182, 
184) with marginal vein punctiform 
pallidipes (p. 51) 

— Ovipositor sheaths brown; forewing (Fig. 189) 
with marginal vein distinctly longer than 
broad pilosus (p. 52) 

41(36) Sensory part of clava extensive and occupying 



14 



42(41) 



43(42) 



44(42) 



45(41) 



46(45) 



an area at least two-fifths as long as clava itself 
giving the clava a strongly transversely or 
obliquely truncate appearance, sutures often 
distinctly oblique (Figs 190, 193, 196, 
202) 42 

Sensory part of clava relatively small and occu- 
pying an area not more than one-fifth as long 
as clava, apex of clava varying from rounded to 
pointed and sutures never oblique (Figs 206, 
208,212,221,227) 45 

Ovipositor not more than about one-third 
longer than mid tibia and not exserted, the 
gonostyli not longer than mid tibial spur and 
yellowish if visible externally 43 

Ovipositor at least twice as long as mid tibia 
and clearly exserted past apex of last tergite, 
the gonostyli about twice as long as mid tibial 
spur and at least partially dark brown 44 

Legs completely yellow; eyes more or less 
naked, with short, sparse setae; frontovertex 
about one-third head width and posterior ocelli 
separated from eye margins by about their own 
diameters valcanus (p. 53) 

Legs with small areas of dark brown near 
apices of femora and bases of tibiae; eyes 
conspicuously hairy, even at low magnifica- 
tions; frontovertex not more than about one- 
quarter head width and posterior ocelli more 
or less touching inner eye margins . urania (p. 
54) 

Antenna (Fig. 196) with all funicle segments, 
except F6, subquadrate or even distinctly 
longer than broad so that pedicel and flagellum 
together are as long as head width; scutellum 
with regular punctate-reticulate sculpture in 
anterior two-thirds or so .. vertumnus (p. 55) 

Antenna (Fig. 202) with all funicle segments, 
except Fl, strongly transverse so that pedicel 
and flagellum together are distinctly shorter 
than head width; scutellum with relatively shal- 
low reticulate sculpture anteriorly .. vesta (p. 
56) 

Forewing (Fig. 207) infuscate below apex of 
venation; tegulae yellow boreas (p. 57) 

Forewing hyaline; tegulae dark brown 46 

Interocellar area very slightly to strongly raised 
above rest of vertex (Figs 210, 215), sculpture 
in this area conspicuously rougher than below 
anterior ocellus, if interocellar area hardly 
raised (Fig. 216) then ovipositor sheaths are 
brown and clearly darker than hind tibia; clava 
pale yellow and at least a little paler than 
funicle 47 

Interocellar area not raised (Figs 219, 224), 
sculpture similar to that below anterior ocellus, 
or if distinctly rougher, then ovipositor sheaths 



D.-W. HUANG AND J.S. NOYES 

honey yellow and more or less concolorous 
with hind tibia; flagellum varying from white to 
brown but all segments concolorous 48 

47(46) Antenna with F6 transverse (Fig. 208); poste- 
rior ocelli separated from occipital margin by 
about 2.5 times their own lengths; scutellum 
convex and dull metallic green slightly mixed 
with coppery towards base clio (p. 58) 

— Antenna with F6 longer than broad (Fig. 212); 
posterior ocelli separated from occipital mar- 
gin by about 1.5 times their own lengths; 
scutellum fairly flat and uniformly bright 
metallic green or blue-green circe (p. 59) 

48(46) Antennae, excluding radicle, completely white 
or very pale yellow clotho (p. 60) 

— Antennae varying from testaceous yellow to 
brown 49 

49(48) All funicle segments longer than broad (Fig. 
221); postmarginal vein of forewing very short, 
less than one-third as long as stigmal vein (Fig. 
222); mandible with a very broad truncation 
with is minutely serrate cybele (p. 61) 

— Antennae at least with Fl-2 quadrate or trans- 
verse (Fig. 227); postmarginal vein at least half 
as long as stigmal; truncate part of mandibles 
not minutely serrate 50 

50(49) Scutellum quite shiny and green to greenish- 
blue particularly in its apical half, anterior part 
slightly coppery; mandibles with two acute 
teeth and a truncation; ovipositor (Fig. 226) 
about 1.5 times as long as mid-tibia and always 
a little exserted ooii (p. 62) 

— Scutellum relatively dull, mostly black with a 
purple tinge, its extreme margins metallic 
green, blue or purple; mandible with one tooth 
and a truncation (or at least second tooth very 
short and strongly obtuse); ovipositor about 
one-third longer than mid-tibia and usually 
hidden 
crassulus (p. 63) 

5 1 (33) Thorax mostly or entirely orange 52 

— Thorax without any orange areas 53 

52(51) Head orange and mesopleura orange 

lucina (p. 63) 

— Head metallic green, mesopleura mostly 
brown 

macula (p. 64) 

53(52) Antenna with Fl transverse and conspicuously 
smaller than other funicle segments which are 
subequal in size and subquadrate (Fig. 242); 
mandible unique with one relatively long tooth 
and an oblique, denticulate truncation (Fig. 
240) 
hera (p. 65) 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



15 



Antenna otherwise, with proximal funicle seg- 
ments subequal, Fl not or conspicuously smaller 
than F2 and frequently much longer than broad 
(Figs 250, 251, 256, 262, 270, 275, 277, 282, 287, 
297, 300, 304, 308, 317-319, 329); mandibles 
either tridentate or with one tooth and a trunca- 
tion (Figs 248, 268, 283, 290, 303, 333, 334) . 54 

54(53) Mandibles apically relatively narrow and with 
three acute teeth, the upper tooth occasionally 
clearly shorter and hardly acute (Figs 248, 268, 
283) 55 

Mandibles relatively broad, with one tooth and 
a broad truncation (Figs 290, 303, 333, 334 and 
as in figs 121 and 139) 62 

55(54) Frontovertex more than one-quarter head 
width (Figs 247, 255,260) 56 

Frontovertex (less than one-quarter head 
width (Fig. 280) 58 

56(55) Antennae (Figs 250, 251) with funicle seg- 
ments subquadrate or longer than broad so 
that clava is about as long as F3-6 together, 
pedicel and flagellum together longer than 
head width; tegulae usually conspicuously yel- 
low or orange basally manii (p. 67) 

— Antennae (Figs 256, 262) with most or all of 
the funicle segments transverse so that clava is 
as long as funicle or nearly so, pedicel and 
flagellum together shorter than head width; 
tegulae entirely brown 57 

57(56) Ovipositor exserted, the exserted part at least 
as long as the mid tibial spur; last tergite 
apically convex, rounded midas (p. 68) 

— Ovipositor not or hardly exserted; last tergite 
with a characteristic V-shaped apical emargin- 
ation mars (p. 69) 

58(55) Antenna with F6 longer than broad; clava with 
sutures parallel, its apex rounded and sensory 
part relatively small and limited to extreme 
apex only neptunus (p. 70) 

— Antenna (Fig. 270) with F6 transverse; clava 
with sutures at least slightly, but distinctly 
convergent, the sensory part enlarged gener- 
ally giving the clava an obliquely truncate 
appearance 59 

59(58) Mesopleuron orange libitina (p. 71) 

— Mesopleuron dark brown or black 60 

60(59) Antenna (Fig. 275) with clava as long as F3-6 
together; forewing (Fig. 276) with a conspicu- 
ous group of setae in basal cell on ventral 
surface close to and running parallel with pos- 
terior wing margin lyaeus (p. 72) 

— Antenna with clava as long as five preceding 
funicle segments together (Figs 277, 282); 
forewing (Fig. 278) without a conspicuous 



group of setae on ventral surface of basal cell 
near posterior wing margin 61 

61(60) Clava brown, contrasting with the yellow 
funicle; last segment of clava occupying about 
two thirds length of clava (Fig. 277) 
lupercus (p. 73) 

Clava and funicle concolorous, pale yellow; 
last segment of clava occupying only slightly 
more than half length of clava (Fig. 282) 
maenas (p. 74) 

62(54) Frontovertex distinctly wider than one-quarter 
head width (Figs 286, 296); scutellum with 
shallow reticulate sculpture which is hardly 
deeper than that on mesoscutum (Figs 288, 
289) 63 

— Frontovertex not wider than one-quarter head 
width (Figs 312, 314, 315); sculpture on scutel- 
lum frequently punctate-reticulate, especially 
in basal half and conspicuously deeper than 
that on mesoscutum (Figs 309, 316) 64 

63(62) Distal funicle segments distinctly longer than 
broad, normally about twice as long as broad 
(Fig. 287); pedicel and flagellum together 
much longer than head width; macropterous 
and brachypterous forms known . segestes (p. 
75) 

— Distal funicle segments hardly longer than 
broad (Fig. 297); pedicel and funicle together 
not or hardly longer than width of head; only 
macropterous forms known .. ferrierei (p. 75) 

64(62) Scutellum basally with relatively shallow sculp- 
ture (Fig. 305) which is not, or hardly, deeper 
than that on mesoscutum 65 

Scutellum, at least in basal half, with deep 
punctate-reticulate or striate-reticulate sculp- 
ture (Fig. 309, 316) which is clearly much 
deeper than that on mesoscutum 66 

65(64) Ovipositor at least slightly exserted, sheaths 
yellow; mandibles with three short teeth and a 
truncation armed with three or four denticles 
(Fig. 303) leander (p. 76) 

Ovipositor hidden, sheaths black or dark 
brown; mandibles with one tooth and a trunca- 
tion armed with numerous very fine, minute 
denticulae musa (p. 77) 

66(64) Basal two-thirds of scutellum with more or less 
striate reticulate sculpture, especially laterally 
(Fig. 309); mesoscutum bright metallic green 
and clothed with conspicuous silvery setae; all 
funicle segments clearly longer than broad 
(Fig. 308) telenomicida (p. 78) 

— Basal half or two-thirds of scutellum with fairly 
regular punctate-reticulate sculpture (Fig. 
316); mesoscutum sometimes bright metallic 
green but setae dark brown, occasionally trans- 



16 

lucent, but never silvery and conspicuous; usu- 
ally one or two funicle segments quadrate or 
transverse (Figs 317-319), but occasionally all 
funicle segments clearly longer than broad 
(Figs 329, 331) 67 

67(66) Funicle with at least two segments quadrate or 
conspicuously smaller than those that follow 
(Figs 317-319); ovipositor at least one-quarter 
longer than mid tibia with sheaths yellow or 
pale brown and about as long as basal tarsal 
segment of hind leg 68 

— Either all funicle segments longer than broad 
and gradually enlarging distad (Figs 329, 331), 
occasionally Fl relatively smaller than F2, or 
ovipositor not longer than mid tibia with 
sheaths dark brown or black and conspicuously 
shorter than basitarsus of hind leg 69 

68(67) Scutellum relatively narrow, not more than 
about 1.3 times as broad as long [longest setae 
on male flagellum at least about 3 times as long 
as diameter of segments (Fig. 328)] 
papilionis (p. 79) 

— Scutellum relatively broad, about 1.5 times as 
long as broad [longest setae on male flagellum 
not more than 1.5 times as long as diameter of 
segments, mostly much shorter (similar to Fig. 
337)] sphingidarum (p. 81) 

69(67) Ovipositor sheaths yellow; tegulae with at least 
base yellow shakespearei (p. 81) 

— Ovipositor sheaths and tegulae completely 
dark brown or black utetheisae (p. 81) 



Review of species 

Ooencyrtus guamensis Fullaway 

(Figs 1-8) 

Ooencyrtus guamensis Fullaway, 1946: 205. 

Holotype , Guam (USNM, examined). 
Schedius garouae Risbec, 1954: 896-899. ?Lecto- 

type , Camerouns (?MNHN, not examined). 

Syn. n. 
Psyllaephagus goarini Risbec, 1958: 98. LECTO- 

TYPE (here designated), Madagascar 

(MNHN, examined). Syn.n. 
Ooencyrtus garouae (Risbec); Annecke & Insley, 

1971:20,39. 

Diagnosis. Mandibles with one tooth and a 
broad truncation; mesoscutum with conspicuous 
silvery hairs; forewing (Fig. 3) with postmarginal 
vein extremely short, basal naked area small and 
open posteriorly; scutellum with deep punctate- 
reticulate sculpture, laterally elongate and 



D.-W. HUANG AND J.S. NOYES 

extreme apex and sides smooth and shiny (Fig. 
2); gaster blackish without yellow or orange 
areas. Female (Length 1.05-1.45 mm): all coxae 
and most part of femora dark brown or black, 
concolorous with mesopleuron; axillae dark 
brown; antennal scape relatively densely hairy; 
clava with a short oblique apical truncation (Figs 
1, 4); F6 subquadrate or a little transverse, other 
funicle segments usually slightly longer than 
broad; frontovertex one-quarter head width; 
hypopygium with anterior margin straight (Fig. 
5). Male (0.79-1.03 mm): legs, including coxae, 
yellow; axillae largely yellow; antennae as in Fig. 
6; frontovertex about two-fifths head width; geni- 
talia as in Fig. 8; fore tibia with a characteristic, 
minutely scaled, elongate depression in its apical 
half (Fig. 7). 

Hosts. This species is a pupal parasitoid of 
Syrphidae (Diptera). It is recorded below from 
Paragus auritus, Asarkira pura, Ischiodon scutel- 
laris; from Africa (see also Prinsloo, 1987) as a 
parasitoid of Ischiodon aegyptius feeding on 
Aphis craccivora, of Paragus borbonicus feeding 
on Dactynotus compositae , of Ischiodon aegyp- 
tius on cotton; and from Hawaii as a parasitoid of 
Allograpta exotica (Beardsley, 1976). The 
records (below) from a coccinellid pupa, from an 
aphid and from caterpillars of Mythimna (=Pseu- 
daletis) unipuncta and Exelastis atomosa (Lepi- 
doptera: Noctuidae) are probably erroneous due 
to incorrect host associations or identifications. 

Distribution. Africa, India, Burma, Thailand, 
Philippines, P. R. China, Malaysia, Indonesia, 
Papua New Guinea, Fiji, Micronesia, Hawaii. 

Material examined. Type material. Holotype 
9 {Ooencyrtus guamensis), GUAM: Piti, ex. 
syrphid puparium, 15. ix. 1936 (O. H. Swezey). 
Paratypes, 1$, Id", same data as holotype 
(USNM). Syntypes of Psyllaephagus goarini Ris- 
bec: 69 labelled 'Psyllaephagus goarini Goarin 
AL366 Lac Alotra', all originally on slides 
(MNHN). Three subsequently remounted on 
cards and the one labelled 'LT9' on underside of 
card is here designated LECTOTYPE. 

Other material. INDIA: 39, Delhi, New 
Delhi, ex. pupa of coccinellid, 1958, CLE. coll. 
No.16082; 39, lcf, Delhi, IARI area, x.1979 (Z. 
Boucek); 1, Uttar Pradesh, Aligarh, 8-10.xi.1979 
(J. S. Noyes); Uttar Pradesh, Dehra Dun, x.1979 
(Z. Boucek); 189, 6cf , Dehra Dun, 2,300ft. ex. 
syrphid pupa, 4.xi.l935/14.xi.l935 (J. A. Gra- 
ham); 29, 2d\ Rajasthan, Udaipur, ex. larva of 
Pseudaletia unipuncta (Haworth), 30. ix. 1972, 
CLE. A6944; 39, 2cf, Rajasthan, Udaipur, ex. 
Pseudaletia unipuncta (Haworth), 30. ix. 1973, 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



17 



CLE. A6602; 1$, Andhra Pradesh, Hyderabad, 
Patancheru, ICRISAT, vii.-ix.1980, M. trap, 
(Bernays & Woodhead); 4$, Maharashtra, 
Poona, ex. moth E. atomosa, CLE. 1474, 
26.viii.1966; 4$, Karnataka, Bangalore, 
19-23. ix. 1979 (J. S. Noyes); 2$, Karnataka, 
Bangalore, xi. 1979 (Z. Boucek); 19, Karnataka, 
Bangalore, ii.1980 (K.D. Ghorpade); 3$, Kar- 
nataka, Bangalore, 916m, ex syrphid pupa, 
28. ii. 1979 (K. D. Ghorpade); Karnataka, Mudig- 
ere, 26.x^.xi.l979 (J. S. Noyes); ld\ Karna- 
taka, 25km. W. of Mudigere, 28.x.-3.xi.l979 (J. 
S. Noyes); 19- Karnataka, Bangalore 916m. ex. 
syrphid pupa, xi. 1971 (K. D. Ghorpade); 1$, 
Karnataka, Bangalore 916m. ex syrphid pupa, 
28. ii. 1973 (K. D. Ghorpade); 3$, Bangalore, ex 
Paragus auritus Stukenberg (det. K.D. Ghor- 
pade, 1986), 14.vii.1986 (T.M. Mushtak Ali); 
29, Tamil Nadu, Coimbatore, 7.xi.l979 (Z. 
Boucek); Tamil Nadu, Coimbatore, 
25.ix-l.x.l979 (J. S. Noyes); 1$, Tamil Nadu, 
Siruvani Forest, 30.ix.1979 (J. S. Noyes); 99, 
5cf, ?Karnataka, Solapur, ex. aphid on saf- 
flower,CIEA20309,iv. 1989. BURMA: 79, lcf, 
Rangoon, 14. i. 1988, CIE A19695. THAILAND: 
29, Chieng Mai, 22.vii.-13.viii. 1984 (D. Jack- 
son); 19, Khao Val National Park, 10-12.U. 1989 
(T. W. Thormin); 29, Huai Kha Khaeng, ii. 1986 
(M.G. Allen). P.R. CHINA: 1$, Hainan, 
coastal vegetation, 19. v. 1983 (Z. Boucek); 39, 
Hainan, Tie Fong Mts., 15. v. 1983 (Z. Boucek); 
19, lo", Hainan,. Tie Fong Mts., 15. v. 1983 (Z. 
Boucek). MALAYSIA: 1$, Sarawak, 4th div. 
Gn. Mulu, RGS Exp., 17.ix.-23.x.l977 (D. Hol- 
lis); 39- 3d", Kuala Lumpur, ex pupa of Asarkiru 
pura, v.1971 (T. H. Chua); PHILIPPINES: 1$, 
Manila, ex syrphid pupa, 2.ix.l959 (C. R. Bal- 
tazar); 19, Manila, ex pupa of Ischiodon scutel- 
laris, 2.ix.l959 (C. R. Baltazar). INDONESIA: 
259, 6d\ Sulawesi Utara, Kotamobagu Danau 
Mooat, 1-6. v. 1985 (J.S. Noyes); 29, Sulawesi, 
Utara, Dumoga-Bone NP, Toraut, 220m. vi.1985 
(A. D. Austin); 99, Sulawesi, Utara, Dumoga- 
Bone NP, Toraut, 220m. 3.V.-6. vi.1985 (J. N. 
Noyes); 69, lcf, Sulawesi, Utara, Dumoga- 
Bone NP, Toraut, 220m. iv.1985 (J. S. Noyes); 
39, Sulawesi, Utara, Dumoga-Bone NP, Toraut, 
MT/YPT. v.1985 (J. S. Noyes); 29, Sulawesi, 
Utara, Dumoga-Bone NP, Toraut, 200m. 
ii.vii.1985. PAPUA NEW GUINEA: 39, 
Bulolo, Mt. Susu, ll.xii.1982 (Z. Boucek); 49, 
Bulolo, 13.xii.1982 (Z. Boucek); 19, Port 
Moresby, 20.xii.1982 (Z. Boucek); 49, 2o\ 
Madang Prov. Laing, 20. vi. 1982 (P. Grootaert). 
FIJI: 19, Viti Levu Korottoongo, hi. 1981 (N. L. 
M. Krauss). Material in BMNH. 
Extralimital material. GHANA: 49, 3d\ with 



aphid on Tecoma radicornis Bignoniaceae, 
17.H. (19)69 em 27. ii. (19)69 (OWR [O.W. Rich- 
ards]); 169- 3d", same data but ex syrphid pupa, 
25. ii em l.iii. (19)69; KENYA: 59, 2o\ Nairobi- 
Chromo, ex syrphid pupa, hi. 1970 (D. Kock); 
39, lcf, Kakamega, ex syrphid pupa on sor- 
ghum, 29.xii.1988 (H. van den Berg); 
P.R.CONGO: Brazzaville, ex syrphid pupa on 
cassava infested with P. manihoti, ? 1 988 (A. 
Biassangama); ZIMBABWE: 279, Chisha- 
washa, various dates v.l980-iv. 1981 (A. Wat- 
sham); ZAMBIA: 1$, Lusaka, 15-24.iv.1980 
(R.A. Beaver); Tanganyika (=TANZANIA): 
239, 9cf, Nachingwea, ex Ischiodon aegyptius 
Wied. feeding on Aphis craccivora em hi. 1954 
(V.F. Eastop); 39, Nachingwea, ex Paragus 
borbonicus Macq. feeding on Dactynotus com- 
posite Theobald (V.F. Eastop); SOUTH 
AFRICA, 179, 4cf. Transvaal, Berberton, ex 
puparium Ischiodon aegyptius on cotton, 1038, 
iii.1921 (G. Ulyett); MADAGASCAR: 29, 
Tulear, Berenty 12 Km NW Amboasarv. 
5-15. v. 1983 (J.S. Noyes, M.C. Day) (compared 
with lectotype of Psyllaephagus goarini Risbec). 
Material in BMNH. 

Comments. We have not examined type mate- 
rial of garouae Risbec, but we have compared 
Oriental material with African material of gar- 
ouae referred to by Prinsloo ( 1987) in his study of 
the Afrotropical species. We are confident that 
garouae is synonymous with guamensis. 

Ooencyrtus guamensis can be separated from 
other species of Ooencyrtus by the following 
combination of characters: both female and 
male with mesoscutum covered by conspicuous 
silvery hairs, antennal scape relatively densely 
hairy (Figs 1, 4, 6), female antennal clava 
enlarged, obliquely truncate at apex (Figs 1.4), 
male fore tibia with a characteristic, minutely 
scaled, elongate depression in its apical half 
(Fig. 7). 



Ooencyrtus acca sp.n. 

(Figs 9-15) 

Diagnosis. Female: all coxae black, femora 
normally at least with black or brown marks 
dorsally; antennae (Fig. 9) with all funicle seg- 
ments transverse; clava conspicuously enlarged, 
with the sutures parallel and with an enlarged 
sensory area, its apex obliquely truncate; fron- 
tovertex about one-fifth head width; posterior 
ocelli nearly touching eye margin; scutellum 
anteriorly with shallow sculpture, the apical half 



18 



D. -W.HUANG AND J.S. NOYES 



smooth (Fig. 12); ovipositor at least slightly 
exserted with gonostyli yellow. 

Female. Length 1.05-1.3 mm (holotype 1.3 
mm). 

Head dark metallic green, ocellar area mixed 
with purple, interantennal prominence mostly 
purple especially dorsally; antennae with radicle, 
scape along dorsal side proximally, dorsal part of 
pedicel, dorsal flagellum and clava testaceous; 
ventral sides of scape, pedicel and funicle seg- 
ments yellow; pronotum, mesoscutum and axil- 
lae dark metallic green or blue mixed with 
purple; anterior half of scutellum with a dark 
purple-brown tinge, duller than mesoscutum, 
posterior half metallic blue mixed with purple; 
mesopleuron and propodeum black-brown to 
blackish; coxae and middle part of femora black- 
brown to blackish; trochanters, both ends of 
femora, tibiae and tarsi yellow, pretarsi brown; 
wings hyaline, venation testaceous; gaster dark 
brown to blackish with some metallic tinge; 
gonostyli yellow. 

Head in ocellar area with fine, raised reticula- 
tions; reticulations on post-ocellar area trans- 
verse; face and genae more shiny than 
frontovertex, sculpture longitudinally elongate; 
ocelli forming an equilateral triangle; posterior 
ocelli separated from occipital margin by about 
1.5 times their own lengths and nearly touching 
eye margins; frontovertex about one-fifth head 
width; eyes conspicuously hairy; occipital margin 
rounded; head in facial view slightly broader 
than high; antennae inserted far below ventral 
eye margins; clypeus with lower margin straight; 
antennae (Fig. 9) with all funicle segments sub- 
equal in length and distinctly transverse; clava 
clearly enlarged, about twice as long as broad, as 
long as F3-6 together; clava with sutures trans- 
verse and with apex obliquely truncate in lateral 
view, sensory area extensive but occupying less 
than one-third length of clava; mandibles with 
one tooth and a broad truncation (Fig. 10). 
Relative measurements (holotype): head width 
48, head height 45, minimum frontovertex width 
9, POL 3.5, OPL 6, OOL 0.2, eye length 30, eye 
width 26, malar space 18, scape length 21, scape 
width 5; other proportions of the antenna as in 
Fig. 9. 

Mesoscutum with somewhat shallow trans- 
versely reticulate sculpture; scutellum (Fig. 12) 
slightly acute posteriorly, its anterior half with 
regular raised reticulate sculpture, posterior half 
smooth and shiny; fore wing with venation and 
distribution of setae basally as in Figs 11 and 13, 
with conspicuous naked area basally which may 
be open or more or less closed posteriorly; linea 



calva open posteriorly; postmarginal vein less 
than half as long as stigmal vein. Relative mea- 
surements (holotype): forewing length 105, 
forewing width 46; hindwing length 72; hindwing 
width 18. 

Gaster at most a little shorter than thorax; 
ovipositor (Fig. 14) slightly exserted, with gono- 
styli nearly one-third as long as ovipositor, and 
second valvifer basally moderately curved; 
hypopygium (Fig. 15) characteristic, with ante- 
rior margin straight and posterior margin 
smoothly curved. Relative measurements 
(paratype): ovipositor length 86, gonostylus 25 
[mid tibia 83]. 

Male. Unknown. 

Variation. There is slight variation in the rela- 
tive width of the frontovertex, setation at the 
base of the forewing and coloration of the thorax 
and mid and hind femora. The femora may be 
largely to almost completely yellow and there 
may be very little blue coloration on the mesos- 
cutum and scutellum. 

Hosts. Unknown. 

Distribution. Brunei, Indonesia. 

Material examined. Type material. Holotype 
9, BRUNEI: Bukit Sulang, Nr. Lamunin, B.M. 
1982-388, 20.viii-10.ix.1982 (N.E. Stork) 
(BMNH). Paratypes, BRUNEI: 3£, same data 
as holotype (BMNH); INDONESIA: 4$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
BMNH Canopy Fog #11, 10.iii.1985; 2$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
iv.1985 (J.S. Noyes) (BMNH). 

Comments. Females of Ooencyrtus acca are 
generally similar to O. guamensis and the Neo- 
tropical species O. syrphidis Noyes. All three 
species have a robust body, antennae with clava 
obliquely truncate at apex and the sensory area 
extensive but less than one-third length of clava, 
forewing with postmarginal vein less than half as 
long as stigmal vein and hypopygium with a 
straight anterior margin (Fig. 15). Ooencyrtus 
acca and syrphidis can both be separated from O. 
guamensis by the absence of silvery setae on the 
mesoscutum, the posterior half of scutellum 
being smooth and the forewing with a realtively 
large conspicuous naked basal area. Ooencyrtus 
acca differs from syrphidis in having F1-F4 trans- 
verse and shallower sculpture on the scutellum 
basally. In syrphidis Fl-4 are at least slightly 
longer than broad and the basal half of the 
scutellum has deep punctate-reticulate sculpture. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



19 



Ooencyrtus phongi Trjapitzin, Myartseva & 
Kostyukov 

(Figs 16-21) 

Ooencyrtus phongi Trjapitzin, Myartseva & 
Kostyukov, 1977: 671-672. Holotype 9, Viet- 
nam (ZISP, not examined). 

Ooencyrtus sp.; Liao et ai, 1987: 176-177. 

Ooencyrtus sp. I; Mehra, 1966. 

Diagnosis. All coxae, femora and gaster black; 
flagellum unicolorous testaceous; scutellum with 
about anterior one-third darker than posterior 
two-thirds; mandibles (Fig. 18) with one tooth 
and a broad truncation, the truncate part with 
numerous denticles; scutellum with deep, regu- 
lar, raised reticulate sculpture covering nearly 
whole surface, only the hind vertical margin 
smooth and shiny; clypeus slightly protuberant 
ventrally. Female (length 1.03-1.71 mm): head 
(Fig. 17) with frontovertex a little more than 
one-quarter head width; posterior ocelli slightly 
but distinctly separate from eyes; antenna (Fig. 
16) with F6 shorter than Fl, proximal 3 funicle 
segments longer than broad, clava less than 3 
times as long as broad, with outer suture oblique 
and apex obliquely truncate; forewing basally as 
in Fig. 19; ovipositor similar to Fig. 335 and 
hypopygium similar to Fig. 36. Male (length 
0.9-1.3 mm): generally similar to female but 
antenna (Fig. 21) with all funicle segments 
clearly longer than broad and setae about twice 
as long as diameter of segments; forewing base 
similar to Fig. 19; genitalia (Fig. 20) with aedea- 
gus about half as long as mid tibia. 

Hosts. Recorded as a parasitoid of the eggs of 
Lepidoptera (Trjapitzin et ai, 1977), but this 
record needs confirmation. The species is 
recorded reliably below from the eggs of Tessa- 
ratoma papillosa, Tessaratoma javanica (see also 
Mehra, 1966) and Pycanum ponderosum (Hemi- 
ptera: Tessaratomidae). It has been recorded as 
an important parasitoid of the eggs of T. papil- 
losa in Thailand (Nanta, 1988), parasitizing more 
than 50% of the eggs of this pest of litchi and 
longan (Dimocarpus longan). 

Distribution. India, Thailand, Vietnam, P.R. 
China, Philippines, Malaysia, Indonesia. 

Material examined. Type material. 

Paratypes, VIETNAM: 8$, Quang-Nin', from 
the eggs of Lepidoptera, 4.vi.l975 (Cam Phong) 
(BMNH). 

Other material: INDIA: 2$, Bihar, Racnhi, 
Namkum, ex eggs of Tessaratoma javanica, 
17.xi.1961 (B. P. Mehra); 4$, Bihar, Ranchi, 



Namkum, ex. eggs of Tessaratoma javanica, 
15.iii.1956 (B.P. Mehra); 1$, Karnataka, 25km 
W. of Mudigere, 28.x-3.xi.1979 (J.S. Noyes); 
Id", Karnataka, Bannerghatta NP, 1979 (Z. 
Boucek& J.S. Noyes); THAILAND: 219, llcf, 
Chiang Mai, ex. eggs of longan bug ( = Tessa- 
ratoma papillosa), iii.1982 (F. Nanta). VIET- 
NAM: 139, 10", Tonkin, Hanoi, from the eggs 
of Tessaratoma papillosa, vi.1937; P.R. CHINA: 
39, Fujian, Kuiqi, ex. eggs of Tessaratoma papil- 
losa, 18.viii.1957 (R. Zhang); 129, 3cf, Fujian, 
Fuzhou, ex. eggs of Tessaratoma papillosa, 
vi. 1991 (Y. Tang); 79, l0\ Hongkong, ex. eggs 
of Tessaratoma papillosa, 16.vi. 1991 (C.Lau); 
239, 3o\ Hainan, Haikou, ex eggs of Tessa- 
ratoma papillosa, 3.vi.l988 (F.D. Bennett & Ren 
Hui); 219, 40", Hainan, Haikou, 25. v. 1983 (Z. 
Boucek); 39. 10". Hainan, Tiefong Mts., 
18. v. 1983 (Z. Boucek); 169, Hainan, Qong- 
zhong, 620m, from litch, 26. vi. 1964 (D.-x. Liao); 
39, Hainan, Qongzhong, Yinggeng, ex. eggs of 
Tessaratoma papillosa, 28. vi. 1964 (D.-x. Liao); 
109. Hainan, Qongzhong, Yinggeng, ex eggs of 
Tessaratoma papillosa, 19. vi. 1964 (D.-x. Liao); 
199, 10d\ Hainan, Xinglong. ex eggs of Tessa- 
ratoma papillosa, 14. vi. 1963 (B. Zhang); 249, 
Hainan, Dangxiang, 7. v. 1983 (Z. Boucek); 
PHILIPPINES: 19, Palawan, Inaguan River 
Valley, 700m, 29.i-2.ii. 1988 (J.H. Martin); 
MALAYSIA: 209. Kuala Lumpur, ex eggs of 
Pycanum ponderosum Stal., 29.vii.1929 (H. Cor- 
bett); INDONESIA: 19, Krakatau, Sertung for- 
est, 15. ix. 1984 (S.G. Compton). Material in 
BMNH, IZAS. 

Comments. We have examined the material 
cited as Ooencyrtus sp. by Liao, et al. (1987) and 
are confident that it belongs to the present spe- 
cies. 

Ooencyrtus acus sp.n. 

(Figs 22-27) 

Echthrodryinus sp., Jadhav & Varma, 1988: 
133. 

Diagnosis. Mandibles more or less tridentate in 
appearance; scutellum with deep reticulate sculp- 
ture and only posterior vertical margin smooth; 
forewing without a conspicuous basal naked 
area. Female: antennae (Fig. 22) with F6 as long 
as or little longer than Fl, the latter subquadrate; 
proximal 4 or 5 funicle segments somewhat 
longer than broad or subquadrate; clava enlarged 
and about 2.5 times as long as broad, with outer 
suture oblique and apex obliquely truncate with 
sensory area occupying nearly half length of 



20 



D. -W.HUANG AND J.S. NOYES 



clava; frontovertex about one-quarter head 
width; posterior ocelli distinctly separated from 
eyes; gaster acute apically and ovipositor slightly 
but distinctly exserted. Male: generally similar to 
female but antennae (Fig. 26) with proximal 
funicle segments slightly longer than broad and 
distal segments quadrate or transverse, the setae 
not longer than diameter of segments; tegulae 
basally yellow. 

Female. Length 0.9-1.3 mm (holotype 1.2 
mm). 

Head blackish, a dark metallic green sheen on 
frontovertex, face metallic green, interantennal 
prominence dorsally purple; antenna with 
radicle, basal dorsal half of pedicel and flagellum 
testaceous, scape and distal half of pedicel yel- 
low; pronotum, mesoscutum and axillae blackish 
with metallic green or blue tinge; tegulae dark 
brown; scutellum metallic blue or green, brighter 
than mesoscutum; mesopleuron and propodeum 
dark brown; coxae and middle part of femora 
dark brown or blackish, ends of femora, tibia and 
tarsi yellow, pretarsi brown, usually at the base 
of tibia with a brown ring; wings hyaline, vena- 
tion brownish; gaster blackish with a slight metal- 
lic green and purple sheen; ovipositor sheath 
blackish. 

Head in ocellar area dull, with fine, raised 
reticulations; face and genae shiny, reticulations 
on them longitudinally elongate; ocelli forming 
an equilateral triangle; posterior ocelli separated 
from occipital margin by about 1.5 times their 
own diameters and conspicuously separated from 
eye margins; frontovertex one fourth head width; 
eyes conspicuously hairy; occipital margin 
slightly rounded; antennae inserted far below the 
ventral eye margin; clypeus with lower margin 
straight; antennae (Fig. 22) with Fl-4 subequal 
in length and subquadrate, and subequal to or 
little shorter than F5 and F6; F6 and sometimes 
F5 somewhat broader than long; clava clearly 
enlarged and about 2.5 times as long as broad, 
about as long as F3-6 together; clava with outer 
suture oblique and apex broadly obliquely trun- 
cate, with sensory area occupying the whole 
ventral surface of last segment; mandibles with 
one tooth and a broad, strongly concave trunca- 
tion which gives the mandibles a more or less 
tridentate appearance (similar to Fig. 268). Rela- 
tive measurements (holotype): head width 44, 
head height 42, minimum frontovertex width 11, 
POL 5, OPL 5, OOL 0.7, eye length 28, eye 
width 23, malar space 15, scape length 19, scape 
width 4; other proportions of the antennae as in 
Fig. 22. 

Mesoscutum with shallow transversely imbri- 



cate sculpture which is clearly shallower than 
sculpture of scutellum; scutellum pointed posteri- 
orly, with regular raised, reticulate sculpture and 
only posterior vertical margin smooth; mesopleu- 
ron with shallow longitudinally elongate sculp- 
ture; forewing with venation and distribution of 
setae basally as in Fig. 23. Relative measure- 
ments (holotype): forewing length 102, forewing 
width 43; hindwing length 65; hindwing width 15. 
Gaster a little longer than thorax plus head; 
ovipositor as in Fig. 24, conspicuously exserted, 
the exserted part about as long as mid tibial spur; 
hypopygium as in Fig. 25. Relative measure- 
ments: ovipositor length 81, gonostylus 20 [mid 
tibia 45]. 

Male. Length about 0.9 mm. 

Similar to female but differs slightly in colora- 
tion, antennal structure, frontovertex width and 
genitalia. Antenna with scape ventrally yellow, 
flagellum testaceous-yellow; legs with mid- 
femora yellow, other femora and coxae brown; 
tegulae basally yellow, brown at apex; antenna as 
in Fig. 26, with proximal funicle segments only 
slightly longer than broad, distal segments quad- 
rate or transverse and setae not longer than 
diameter of segments; frontovertex two-fifths to 
one-third head width; posterior ocellus separated 
from occipital margin by less than its length (3: 
4); gaster conspicuously shorter than head and 
thorax together; genitalia as in Fig. 27 with 
aedeagus about as long as mid tibia. 

Variation. Very little in material available 
apart from that discussed above under coloration 
and antennal structure. 

Hosts. Jadhav & Varma (1988) recorded this 
species (as Echthrodryinus sp.) as a gregarious 
hyperparasitoid of the prepupae or pupae of 
Epiricania melanoleuca (Lepidoptera: Epipy- 
ropidae), an important ectoparasitoid of Pyrilla 
perpusilla (Homoptera: Lophopidae) which is a 
common pest of sugarcane in India and else- 
where. 

Distribution. India. 

Material examined. Type material. Holotype 
$, INDIA: Maharashtra, Dadh, ex cocoon of 
Epricania melanoleuca, 31.x. 1991 (R.B. Jadhav) 
(BMNH). Paratypes, 11$, ld\ same data as 
holotype; INDIA: 5$, Maharashtra, Chande- 
gaon, ex cocoon of Epipyrops melanoleuca, CIE 
A15767, 25.x. 1985 (R.B. Jadhav); 4$, ?Maha- 
rashtra, Satral, ex cocoon of Epipyrops melano- 
leuca, CIE A15767, 2.L1984 (R.B. Jadhav), 
Echthrodryinus sp. det. B.R. Subba Rao, 1984; 
3$, Maharashtra, Ravalgaon, ex pupa of Epipy- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



21 



rops sp., CIE A15395, 5.xi.l982, Echthrodryinas 
sp., det. B.R. Subba Rao, 1983; 6$ (one on 
slide), ld\ Maharashtra, Sade, ex. pupa of 
Epipyrops melanoleuca, CIE A14657, 13.ixl982 
(MSG), Echthrodryinus sp. det. B.R. Subba 
Rao, 1983; 3$, ld\ Gujarat, Navsari, ex. pupa 
of Epipyrops melanoleuca, CIE A15356, 9. i. 1983 
(Z. Patel), Echthrodryinus sp. det. B.R. Subba 
Rao, 1983; 39, Uttar Pradesh, Lucknow, ex. 
cocoon of Epipyrops, CIE A11011, x.1976 (N. 
Nigam), ? Echthrodryinus sp., 1977; 3$, Orissa, 
Sankumari, ex. pupa of Epiricania melanoleuca, 
CIE A19437, xi.1987. Material in BMNH. 

Comments. Females of Ooencyrtus acus share 
the following common characters with O. phongi 
and O. lucens: scutellum with fairly deep, regular 
raised reticulations, with only posterior vertical 
margin smooth; clava enlarged, with outer suture 
oblique and apex obliquely truncate and 
enlarged sensory area that is nearly half as long 
as clava. Ooencyrtus acus can be separated from 
phongi and lucens by the tridentate appearance 
of the mandibles, acute apex of the gaster, the 
ovipositor distinctly exserted, the forewing 
basally without conspicuous naked area (com- 
pare Figs 23, 19 and 35) and the second valvifers 
basally being relatively narrow and more strongly 
curved. In phongi and lucens the mandibles have 
a broad, straight truncation, the gaster is apically 
truncate, the ovipositor is not exserted, and the 
base of the forewing has a more or less conspicu- 
ous naked area. 

Ooencyrtus caurus sp.n. 

(Figs 28-33) 

Diagnosis. Mandibles with one very small 
tooth and a broad truncation; legs with coxae and 
femora black. Female: body black, dorsum of 
thorax bright metallic blue or green mixed with 
purple; gaster shorter than thorax; scutellum 
with posterior one-third smooth, anteriorly with 
punctate-reticulate sculpture; clava with outer 
suture oblique, its apex obliquely truncate with 
sensory part nearly half as long as clava; F1-F3 
small, transverse, conspicuously smaller than dis- 
tal funicle segments (Fig. 28); vertex one-fifth 
head width; basal cell of forewing posteriorly 
without hairs on ventral side, basal naked area 
large, nearly closed posteriorly (Fig. 29), post- 
marginal vein very short, about one-third as long 
as stigmal vein. Male: generally much duller than 
female, but with scutellum bright metallic green; 
antenna with longest setae on funicle about 3 
times as long as diameter of any segment; Fl 
shortest, slightly longer than broad and as long as 



pedicel (Fig. 32), F2-6 each about 1.5 times as 
long as broad. 

Female. Length, including ovipositor: 0.65- 
0.95 mm (holotype 0.82 mm). 

Head black, dull purplish mixed with green 
especially on face and genae, shining green 
between ocelli and occipital margin; antennae 
yellow, only pedicel proximally somewhat 
brownish; pronotum blackish; mesoscutum with 
strong dark blue or purple sheen; axillae black- 
ish; scutellum with anterior three-fifths green, 
posterior two-fifths metallic blue; mesopleuron 
and propodeum dark brown or black; legs with 
coxae and femora excluding tip black, concolor- 
ous with mesopleuron; wings hyaline, venation 
testaceous-yellow; gaster black dorsally with 
strong basal metallic blue or green sheen. 

Head with shallow, regular hexagonally reticu- 
late sculpture on frontovertex, lower parts of 
face laterally and genae with elongate reticula- 
tions; ocelli forming an angle of about 45°; poste- 
rior ocelli separated from occipital margin by 
about 1.5 times their own length and almost 
touching eye margins; eye without conspicuous 
setae; occipital margin sharp, but not carinate; 
antennae inserted far bellow the ventral eye 
margin; antennae (Fig. 28) with pedicel plus 
flagellum conspicuously shorter than head width; 
Fl-3 conspicuously smaller than F4-6 which are 
subquadrate; clava as long as F3-6 together and 
with outer suture strongly oblique and its apex 
obliquely truncate with sensory area nearly half 
as long as clava; mandibles with one very small 
outer tooth and a broad truncation armed with 
minute denticles. Relative measurements (holo- 
type): head width 46, head height 40, minimum 
frontovertex width 9, POL 5, OOL 0.5, eye 
length 27, eye width 25, malar space 15, scape 
length 17, scape width 4; other proportions of 
antennae as in Fig. 28. 

Mesoscutum about 1.2 times as long as scutel- 
lum and about 1.7 times as broad as long, with 
shallow transverse imbricate sculpture; scutellum 
anteriorly with deep, regular reticulate sculpture 
its posterior one-third smooth; forewing with 
venation and basal setation as in Fig. 29. Relative 
measurements (holotype): forewing length 93, 
forewing width 38; hindwing length 62, hindwing 
width 14. 

Gaster in specimens at hand extremely com- 
pact, less than half thorax length (although in 
critical point dried material the gaster may be 
much more distended); ovipositor hidden; ovi- 
positor (Fig. 30) with second valvifers basally 
moderately curved and gonostyli short; hypopy- 
gium as in Fig. 31. Relative measurements 



22 



D. -W.HUANG AND J.S. NOYES 



(paratype): ovipositor length 40, gonostylus 7 
[mid tibia 42]. 

Male. Length about 0.65 mm. 

Differs from female in coloration and structure 
of the antennae, relative width of frontovertex, 
duller coloration, and structure of genitalia. 
Antennae (Fig. 32) with radicle, scape and 
pedicel brown, flagellum testaceous; mesoscu- 
tum dull with a slight brassy and purplish sheen; 
frontovertex two-fifths head width; ocelli form- 
ing a equilateral triangle; posterior ocelli sepa- 
rated from occipital margin by their own lengths 
and conspicuously separated from eyes; genitalia 
as in Fig. 33, with aedeagus about 0.75 times as 
long as mid tibia. 

Variation. In the female the frontovertex var- 
ies from about one-fifth to one-sixth head width; 
the flagellum may be unicolorous yellow or with 
clava conspicuously darker and brownish. 

Hosts. Recorded below from the eggs of 
Amblypelta sp. (Homoptera; Coreidae) on Abel- 
moschus manihot and from the eggs of 
Amblypelta lutescens, a pest of banana in Queen- 
sland, Australia (H.A.C. Fay, pers. comm.). 

Distribution. Papua New Guinea, Australia. 

Material examined. Type material. Holotype 
J, PAPUA NEW GUINEA: Central Province, 
Laloki, ex Amblypelta eggs on Abelmoschus 
manihot, 24. vi. 1983 (F. Dori). Paratypes, 10$, 
2cf (1 on slide), same data as holotype; 5$, 
Laloki Rs. Stn, ex eggs of Amblypelta lutescens 
papuensis, CIE A20325, 24.viii.1989 (F. Dori); 
AUSTRALIA: 4$, 3cf, Queensland, Mareeba, 
ex eggs of Amblypelta lutescens lutescens (Dis- 
tant) on banana, 6.x. 1992 (H.A.C. Fay). Mate- 
rial in BMNH. 

Comments. Ooencyrtus caurus is very close to 
lucens, females of both species having an 
obliquely truncate clava, relatively short anten- 
nal segments, similar mandibular stucture, gener- 
ally similar habitus and body coloration. O. 
caurus can be separated from lucens in having the 
posterior one-third of the scutellum completely 
smooth and polished whereas in lucens the 
scutellum is almost entirely reticulate. See also 
comments under lucens (below). 

Ooencyrtus lucens sp.n. 

(Figs 34-36) 

Diagnosis. Female: all coxae and femora dark, 
concolorous with mesopleuron; scutellum wholly 
bright green-blue, with moderately deep- 



punctate sculpture over most of its surface and 
only its extreme posterior margin smooth; anten- 
nae (Fig. 34) with flagellum usually with funicle 
testaceous-yellow and clava brownish; F6 longer 
than Fl, Fl-3 transverse; clava with outer suture 
oblique and apex obliquely truncate and with 
sensory area about half as long as clava; fron- 
tovertex about one-fifth head width. 

Female. Length 0.70-1.15 mm (holotype 0.95 
mm). 

Very similar to caurus but differing in the 
deeper, more extensive reticulate sculpture of 
the scutellum (see diagnosis). Antennae as in 
Fig. 34; fore wing base as in Fig. 35; hypopygium 
as in Fig. 36; ovipositor similar to Fig. 335. 
Relative measurements (holotype): head width 
46, head height 44, minimum frontovertex width 
7.5, POL 3.5, OPL 5, OOL 0.3, eye length 31, 
eye width 24, malar space 18, scape length 19, 
scape width 4.5; forewing length 96, forewing 
width 40; hindwing length 65, hindwing width 16; 
(paratype): mid tibia 90, ovipositor length 87, 
mid tibia 90, gonostylus 15. 

Male. Unknown. 

Hosts. Recorded below from the eggs of 
Achaea janata (Lepidoptera: Noctuidae) and 
from an unidentified pentatomid (Hemiptera: 
Pentatomidae). 

Distribution. India, Brunei, Malaysia, Philip- 
pines. 

Material examined. Type material. Holotype 
$, BRUNEI: Bukit Sulang, Nr. Lamunin, B.M. 
1982-338, 20.viii-10.ix.1982 (N.E. Stork) 
(BMNH). Paratypes, INDIA: 1$, Karnataka, 
25km W. of Mudigere, 28.x-3.xi. 1979 (J.S. 
Noyes); INDIA: 3$, Andaman Islands, Port 
Blair, ex Achaea janata, 3.vii.l992 (K. Veenaku- 
mari); MALAYSIA: 1$, Sabah, Longkong, ex 
Pentatomid, CIE A13458, 16. ix. 1981 (G.J. Lim); 
BRUNEI: 7$, same data as holotype; 1$, 
PHILIPPINES: Leyte, Baybay, ix.1980 (R. 
Vane-Wright). 

Comments. Ooencyrtus lucens is most similar to 
phongi and caurus, all three species having man- 
dibles with one small tooth and a broad straight 
truncation; coxae and most part of femora dark, 
concolorous with mesopleuron; scutellum deeply 
reticulate at least in anterior half or so; clava with 
outer suture oblique and its apex obliquely trun- 
cate with sensory part very extensive, about half 
as long as clava. Females of both lucens and 
caurus can be separated from phongi by the lack 
of silvery setae on the meoscutum, F6 being 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



23 



longer than Fl (in phongi F6 is shorter than Fl); 
flagellum at least partially yellowish (unicolorous 
dark brown in phongi) and the scutellum wholly 
brightly metallic (in phongi the anterior part of 
the scutellum is relatively dull). See also com- 
ments under caurus. 

Ooencyrtus minerva sp.n. 

(Figs 37, 38) 

Diagnosis. Female: body, excluding legs, gen- 
erally blackish with a weak metallic green or 
bluish sheen on the frontovertex and dorsum of 
thorax and gaster; funicle segments dark testa- 
ceous, clava yellow; all coxae and femora, except 
apices, black; antennae (Fig. 37) with flagellum 
filiform and all funicle segments longer than 
broad, clava not broader than funicle segments 
and with sensory area at apex only; mandibles 
with one small tooth and a very broad straight 
truncation; scutellum in anterior three-fifths with 
regular, raised punctate-reticulate sculpture, pos- 
terior two-fifths smooth and shiny; forewing (Fig. 
38) without distinct basal naked area, linea calva 
closed posteriorly. 

Female. Length 1.35 mm. 

Head on frontovertex with a weak metallic 
green sheen, face with a stronger metallic blue- 
green reflection; interantennal prominence cop- 
pery purple, lower face and genae with a slight 
purple lustre and less shiny; antennae with 
radicle black, dorsal sides of scape black and rest 
of scape testaceous; pedicel and funicle segments 
dark testaceous, clava yellow; pronotum, mesos- 
cutum and axillae blackish with a weak metallic 
blue tinge; scutellum with anterior three-fifths 
dark green and relatively dull, posterior two- 
fifths blue, shiny, much brighter than anterior 
part; mesopleuron and propodeum blackish; 
coxae, femora except apices blackish, trochant- 
ers, apex of femora, tibia and tarsi testaceous to 
yellow; wings hyaline, venation testaceous; 
gaster blackish with a metallic green and purple 
tinge; ovipositor sheaths blackish. 

Head in ocellar area with fine, raised reticula- 
tions; reticulations on genae longitudinally elon- 
gate; ocelli forming a 45° angle, posterior ocelli 
separated from occipital margin by about 1.5 
times their own lengths and almost touching eye 
margins; frontovertex about one-sixth head 
width; eyes naked, without conspicuous setae; 
occipital margin slightly rounded; antennae 
inserted just below the ventral eye margins, 
toruli separated from mouth margin by little 
more than their own lengths; antennae (Fig. 37) 
with all funicle segments conspicuously longer 



than broad, clava a little shorter than F4-6 
together, with sensory part limited at apex only; 
mandibles with one small tooth and a very broad 
straight truncation. Relative measurements: 
head width 53, head height 50, minimum fron- 
tovertex width 9, POL 4.5, OPL 4.5, OOL 0.5, 
eye length 37, eye width 30, malar space 20, 
scape length 25, scape width 5; other proportions 
of the antenna as in Fig. 37. 

Mesoscutum with shallow, transverse 
imbricate-reticulate sculpture; scutellum in basal 
three-fifths with deep, regular, punctate- 
reticulate sculpture, posterior two-fifths smooth; 
fore wing with venation and distribution of setae 
basally as in Fig. 38. Relative measurements: 
forewing length 123, forewing width 50; hind- 
wing length 82, hindwing width 20. 

Gaster shorter than thorax; ovipositor not 
exserted, the gonostyli very short, only very 
slightly longer than half length of mid tibial spur. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Malaysia. 

Material examined. Type material. Holotype 
9, MALAYSIA: GentingTea Est., vii-viii. 1985, 
2000m (W. Budenberg) [right wings and antenna 
mounted on slide] (BMNH). 

Comments. Ooencyrtus minerva and ceres are 
similar in that both species have a bicolorous 
flagellum with the funicle dark testaceous and 
clava yellow, all funicle segments longer than 
broad, scutellum with punctate sculpture anteri- 
orly and smooth posteriorly and the forewing 
with the linea calva closed posteriorly. The two 
species can be separated on the relative width of 
the frontovertex (minerva one-sixth and ceres 
one quarter head width), relative length of the 
pedicel and flagellum together (minerva 1.4 
times and ceres 1.2 times head width), relative 
proportions of the antennal segments (minerva 
F6 shorter than Fl and ceres Fl shorter than F6) 
and coloration of scutellum (minerva anteriorly 
dark green and dull but in ceres the scutellum is 
entirely green-blue). 

Ooencyrtus icarus sp.n. 

(Figs 39-40) 

Diagnosis. Female: antennae with radicle and 
proximal two-thirds of scape brownish, flagellum 
testaceous, all coxae and femora including mid- 
femora dark brown of black; mandibles with one 
very small tooth and a broad truncation; flagel- 
lum filiform with all funicle segments longer than 



24 



D. -W.HUANG AND J.S. NOYES 



broad, clava not broader than funicle segments, 
its apex pointed and with a small sensory area at 
apex only; scutellum with regular fine punctate- 
reticulate sculpture, in posterior fifth or so shal- 
lower and with posterior margin smooth, apex 
moderately pointed; forewing (Fig. 40) without 
conspicuous naked area basally; linea calva 
closed posteriorly. 

Female. Length about 0.95 mm. 

Head blackish with a metallic green sheen, 
interantennal prominence purple; antennae with 
radicle and proximal two-thirds of scape black- 
ish, distal one-third of scape testaceous, pedicel 
proximally brownish and distally testaceous, fla- 
gellum testaceous; pronotum, mesoscutum, axil- 
lae and anterior one-third of scutellum blackish 
with weak metallic green, coppery or brassy 
tinge; posterior two-thirds of scutellum metallic 
blue, much brighter than anterior part which is 
relatively dull and purplish; mesopleuron and 
propodeum blackish; coxae and femora except 
the extreme ends concolorous with mesopleuron, 
both ends of femora yellow; wings hyaline, vena- 
tion testaceous; gaster blackish with a metallic 
green and purple tinge; ovipositor sheath black. 

Head in ocellar area with fine, raised reticulate 
sculpture; reticulations on genae longitudinally 
elongate; ocelli forming an acute triangle (about 
40°), the posterior ocelli separated from occipital 
margin by about their own lengths and almost 
touching eye margins; frontovertex a little more 
than one-fifth head width; eyes clothed in short, 
inconspicuous translucent setae; occipital margin 
acute; antennae inserted just below the ventral 
eye margin; toruli separated from mouth margin 
by a little less than their own lengths; antennae 
(similar to Fig. 32) with pedicel plus flagellum as 
long as head width; all funicle segments con- 
spicuously longer than broad, subequal in length; 
clava about as long as preceding three segments 
together, with sensory area very small and incon- 
spicuous; mandibles (Fig. 41) with one obtuse 
small tooth and a broad truncation bearing 
minute denticulae. Relative measurements 
(holotype): head width 46, head height 41, mini- 
mum frontovertex width 10, POL 5, OPL 2.5, 
OOL 0.2, eye length 28, eye width 25, scape 
length 17, scape width 4; other proportions of the 
antenna similar to Fig. 32. 

Thorax with shallow, transverse imbricate- 
reticulate sculpture on mesoscutum; scutellum 
with deeper, regular, finely reticulate sculpture 
becoming shallower in apical one-fifth or so with 
apex moderately pointed and nearly smooth; 
fore wing with venation and distribution of setae 
basally as in Fig. 40, without a conspicuous 



naked area, linea calva closed posteriorly, post- 
marginal vein half as long as stigmal vein. Rela- 
tive measurements (paratype): forewing length 
99, forewing width 41; hindwing length 70; hind- 
wing width 16. 

Gaster with ovipositor hidden; ovipositor simi- 
lar to Fig. 335, with gonostyli about one-fifth as 
long as ovipositor; hypopygium as in Fig. 39. 
Relative measurements: ovipositor length 107, 
gonostylus 20 [mid tibia 95]. 

Male. Unknown. 

Hosts. Recorded below from the eggs of Dasy- 
nus kalshoveni (Hemiptera: Coreidae). 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
9, INDONESIA: Java, Semarang, ex eggs of 
Dasynus kalshoveni, iv.1937 (J.S. Phillips). 
Paratype, 19 (on slide) same data as holotype. 
All material in BMNH. 

Comments. Females of Ooencyrtus icarus are 
similar to those of iris, both species having the 
mandible with a broad, minutely denticulate 
truncation, relatively long funicle segments, uni- 
colorous flagellum and forewing with a closed 
linea clava. However, in O. icarus the mid 
femora are blackish (yellow in iris) and the 
scutellum has finely reticulate (almost punctate) 
sculpture over most of its surface (anterior half 
shallow and irregular and posterior half smooth 
and shiny in iris). Superficially both species also 
resemble caurus and lucens but the antennae are 
more filiform and the clava lacks the obliquely 
truncate apex with an extensive sensory area. 

Ooencyrtus adonis sp.n. 

(Figs 42-44) 

Diagnosis. Female: flagellum unicolorous; legs 
with coxae and femora dark brown, concolorous 
with mesopleuron, tibiae each proximally dark 
brown or with a dark ring; gaster totally brown; 
mandibles with four teeth (similar to Fig. 46); 
scutellum with extremely shallow sculpture in 
anterior half and with posterior half smooth; 
antennae (Fig. 42) with Fl-4 relatively small, 
transverse or quadrate and much smaller than 
either F5 or F6 which are subequal in size; clava 
not conspicuously broader than funicle segments, 
apically more or less rounded and with only a 
small sensory area at apex; frontovertex about 
two-fifths head width; posterior ocelli distinctly 
separated from eyes; ovipositor far exserted, 
exserted part from half to one-third gaster 
length. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



25 



Female. Length, excluding ovipositor, 0.5-0.68 
mm (holotype 0.68 mm). 

Frontovertex and genae dark brown with weak 
metallic green and purple tinge, lower face 
including interantennal prominence slightly paler 
brown; antennae with radicle, dorsal sides of 
scape and pedicel brownish; ventral sides of 
scape and pedicel and flagellum testaceous- 
yellow; pronotum, mesoscutum, axillae and ante- 
rior half of scutellum dark brown with a weak 
metallic green or purple sheen; scutellum poste- 
riorly shiny blue-green shiny; mesopleuron 
brown; propodeum dark brown; legs with coxae 
and femora brown, tibiae proximally dark brown 
or with a dark ring; wings hyaline, venation 
testaceous; gaster brown; ovipositor sheaths 
brown. 

Head in ocellar area with fine, raised reticula- 
tions; sculpture on frons, lower face and genae 
longitudinally elongate; ocelli forming a slightly 
obtuse angle, posterior ocelli separated from 
occipital margin by at most their own lengths and 
distinctly separated from eye margins; frontover- 
tex about two-fifths head width (Fig. 43); eyes 
without conspicuous setae; occipital margin more 
or less rounded; antennae inserted slightly below 
ventral eye margin (Fig. 43), toruli separated 
from lower margin of face by their own length or 
less; antennae (Fig. 42) with Fl-4 relatively 
small, transverse or quadrate and much smaller 
than either F5 or F6 which are subequal in size; 
clava as long as F3-6 together, not conspicuously 
broader than funicle segments, apically more or 
less rounded and with only a small sensory area 
at apex; mandibles (similar to Fig. 46) with four 
teeth. Relative measurements (holotype): head 
width 24, head height 21, minimum frontovertex 
width 10, POL 5, OPL 1.5, OOL 1, eye length 
14, eye width 10.5, malar space 9, scape length 
10, scape width 2; other proportions of the 
antennae as in Fig. 42. 

Mesoscutum with transverse imbricate- 
reticulate sculpture which is deeper than sculp- 
ture of scutellum; scutellum with very shallow 
longitudinally elongate polygonal sculpture in 
anterior half, the posterior part smooth; fore- 
wing with venation and distribution of setae 
basally as in Fig. 44. Relative measurements 
(holotype): forewing length 56, forewing width 
25; hindwing length 41, hindwing width 9. 

Gaster, including ovipositor, about as long as 
thorax or shorter; ovipositor (similar to Fig. 49) 
far exserted, the exserted part about one-third to 
half gaster length, or about half length of mid- 
tibia; hypopygium similar to Fig. 50. Relative 
measurements (paratype): ovipositor length 41, 
gonostylus 11.5 [mid tibia 19]. 



Male. Unknown. 

Hosts. Unknown. 

Distribution. Nepal. 

Material examined. Type material. Holotype 
$, NEPAL: Bardia, 10-24.iii.1983 (M.G. 
Allen). Paratypes, 39, same data as holotype. 
All material in BMNH. 

Comments. Ooencyrtus adonis is probably clos- 
est to aeneas, both species having brown coxae 
and femora, relatively wide frontovertex, shallow 
sculpture on scutellum, ovipositor well exserted 
and mandibles with four teeth. O. adonis can be 
separated from aeneas by the structure of the 
antennae and relative length of the exserted part 
of the ovipositor. In aeneas the funicle segments 
are all subquadrate and subequal in size, except 
for Fl which is conspicuously transverse and the 
exserted part of the ovipositor is about one- 
quarter as long as the gaster. 

Ooencyrtus aeneas sp.n. 

(Figs 45-50) 

Diagnosis. Coxae and femora dark brown, 
tibiae mostly dark brown; mandibles with four 
teeth (Fig. 46); antennae (Fig. 47) with Fl small 
and transverse, much smaller than F2-6 which 
are slightly longer than broad and subequal; 
clava apically rounded with sensory area on apex 
only; head with frontovertex about one-third 
head width; posterior ocelli distinctly separated 
from eyes; scutellum anteriorly with shallow 
sculpture and posteriorly smooth and shiny; ovi- 
positor exserted, exserted part about one-quarter 
as long as gaster. 

Female. Length, excluding ovipositor, 0.83 
mm. 

Head dark brown with a slight brassy sheen on 
frontovertex and greenish in scrobal area; anten- 
nae with radicle, dorsal sides of scape and pedicel 
brown; ventral sides of scape and pedicel and 
flagellum testaceous; pronotum, mesoscutum 
and axillae dark brown or blackish with a slight 
purple lustre; scutellum with anterior three-fifths 
blackish with a purple sheen, posterior two-fifths 
metallic green or blue; mesopleuron brown; pro- 
podeum blackish; legs with coxae and femora 
brown, tibiae with basal three-fifths dark brown; 
wings hyaline, venation testaceous; gaster 
brown; ovipositor sheath brown. 

Head in ocellar area with fine, raised reticulate 
sculpture; sculpture on lower face and genae 
longitudinally elongate; ocelli forming a right 
angle, posterior ocelli about equidistant from 



26 



D. -W.HUANG AND J.S. NOYES 



occipital and eye margins; frontovertex one-third 
head width; eyes without conspicuous setae; 
occipital margin more or less rounded; antennae 
inserted slightly below ventral eye margin (Fig. 
45); toruli separated from mouth margin their 
own lengths; antennae (Fig. 47) with Fl trans- 
verse and clearly much smaller than any of the 
following segments which are wider and at least 
slightly longer than broad; clava with a small 
sensory area at apex only; mandibles (Fig. 46) 
with four teeth. Relative measurements (holo- 
type): head width 29, head height 25, minimum 
frontovertex width 10, POL 4.5, OPL 1.5, OOL 
1, eye length 17, eye width 13, malar space 11, 
scape length 11, scape width 2.5; other propor- 
tions of the antennae as in Fig. 47. 

Mesoscutum with transverse imbricate sculp- 
ture which is clearly deeper than that on scutel- 
lum; sculpture on anterior three-fifths of 
scutellum shallow and more or less longitudinally 
elongate or polygonal espcially medially, poste- 
rior two-fifths smooth and shiny; fore wing with 
venation and distribution of setae basally as in 
Fig. 48. Relative measurements (holotype): 
forewing length 71, forewing width 28; hindwing 
length 45; hindwing width 10. 

Gaster, excluding ovipositor, about as long as 
thorax; ovipositor (Fig. 49) exserted, the 
exserted part about one-quarter as long as gaster; 
hypopygium as in Fig. 50. Relative measure- 
ments: ovipositor length 107, gonostylus 30 [mid 
tibia 55]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
9, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 200m, BMNH Canopy Fog 
#13, ll.vii.1985. Paratype, 1$, BMNH Canopy 
Fog #11, 10. hi. 1985. Material in BMNH. 

Comments. Ooencyrtus aeneas is close to adonis 
(see comments under adonis above). 

Ooencyrtus phoebi nom. nov. 

Ooencyrtus major Ferriere, 1931: 285. LECTO- 
TYPE $ (here designated), Indonesia 
(BMNH, examined). 

Diagnosis. Females (length 1.4-1.5 mm): body 
including gaster black, weakly metallic green or 
purple, scutellum in apical half or so more 
strongly metallic blue-green; legs with coxae and 
fore femora dark brown, mid femora mostly 
yellow, hind femora and tibiae mostly dark 



brown, fore and mid tibiae mostly yellow; man- 
dibles with one minute tooth and a very broad 
truncation; scape slender; flagellum filiform; all 
funicle segments distinctly longer than broad; 
clava hardly wider than funicle, with sutures 
parallel and with sensory area at extreme apex 
only; eyes with conspicuous short setae; fron- 
tovertex at most one-quarter head width; scutel- 
lum with reticulate sculpture similar to that on 
mesoscutum, becoming shallow and shiny 
towards apex which is slightly pointed; forewing 
with a large basal naked area closed posteriorly; 
ovipositor slightly exserted, yellow. Males: none 
examined, but according to the original descrip- 
tion all funicle segments are longer than broad 
and clothed with short setae. 

Hosts. Reared from the eggs of Attacus atlas 
(Lepidoptera: Saturniidae). According to Fer- 
riere (1931) other egg parasitoids were also bred 
from the eggs of the same host, including Anasta- 
tus menzeli Ferriere, (Eupelmidae), Agiommatus 
attaci Ferriere (Pteromalidae) and Tetrastichus 
sp. (Eulophidae). 

Distribution. Indonesia. 

Material examined. Type material. Lecto- 
type 9, INDONESIA: Java, Buitenzorg, ex eggs 
of Attacus atlas F., vii.1925 (R. Menzel), 
B.M.TYPE HYM. 5. 1,073, originally with a red 
'type' label and Ferriere's handwriting '9 type'. 
Paralectotypes: 39, same data as the lectotype 
but each with a yellow 'Co-type' label (BMNH). 

Comments. This species seems to be closest to 
Ooencyrtus kuvanae (Howard) both species 
being relatively large, only weakly metallic, the 
legs extensively dark brown, the flagellum fili- 
form with all funicle segments longer than broad, 
the sensory area of the clava limited to the 
extreme apex, the frontovertex at most one- 
quarter head width, and the forewing with a large 
basal naked area, closed posteriorly. O. phoebi 
can be separated from kuvanae by the conspicu- 
ously hairy eyes, paler leg coloration and shal- 
lower sculpture of the scutellum (see key). 

The species name major is preoccupied in 
Ooencyrtus by Ooencyrtus major (Perkins, 1906). 

Ooencyrtus kuvanae (Howard) 

(Figs 51-53) 

Schedius kuvanae Howard, 1910: 3-5. Holotype , 

Japan (USNM) [not examined]. 
Ooencyrtus kuwanael (Howard); Mercet, 1926: 

48-50. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



27 



Ooencyrtus kuwanai (Howard); Peck, 1951: 496. 
Invalid emendation of kuvanae. 

Other citations. For a comprehensive refer- 
ence list see Peck, 1963: 428-431; Brown, 1984 
and Trjapitzin, 1989: 207-208. 

Diagnosis. Female (length 0.87-1.35 mm): 
body including gaster black or brown with a weak 
metallic green, blue or purple reflections on head 
and pronotum, mesoscutum with a relatively 
strong bluish sheen; scutellum medially in ante- 
rior half coppery purple, remainder metallic 
green, blue or purple; legs with coxae and at least 
proximal part of femora concolorous with 
mesopleuron, dark brown; ovipositor sheaths 
yellow; eyes without conspicuous setae; antennae 
(Fig. 51) with flagellum filiform, unicolorous; all 
of funicle segments distinctly longer than broad; 
clava with sensory area at extreme apex only; 
frontovertex about one-quarter head width; man- 
dibles with a small outer teeth and a broad inner 
truncation; forewing (Fig. 53) with a small basal 
naked area which is closed posteriorly; scutellum 
almost entirely with fine reticulate sculpture 
which is conspicuously deeper than that on 
mesoscutum, only its vertical sides and apex 
smooth and polished; ovipositor very slightly 
exserted; hypopygium as in Fig. 52. Male (length 
0.71-0.95 mm): generally similar to female but 
scutellum relatively dull; antennae with flagellum 
testaceous-yellow, all funicle segments about 2 
times as long as broad and with longest setae 
about as long as diameter of segments; frontover- 
tex about one-third head width. 

Hosts. Ooencyrtus kuvanae (Howard) is known 
primarily as an egg parasite of the gypsy moth, 
Lymantria dispar L. and may play an important 
role as a biological control agent of this well- 
known pest species (Brown, 1984). It has also 
been recorded from the eggs of lasiocampids 
including Lymantria fumida (Trjapitzin, 1989), 
Dendrolimus spectabilis and Malacosoma neus- 
tria (below), saturniids including Eriogyna pyre- 
torum (Koidzumi & Shobata, 1940) and other 
lymantriids including Stilpnotia salicis (below) 
and Nygmia phaeorrhoea Donovan. In the labo- 
ratory, it has been reared successfully on the eggs 
of Euproctis chrysorrhoea, Hemerocampa leu- 
costigma, Hemerocampa definata (Lymantri- 
idae), Hemileuca maia, H. oliviae, Callosamia 
promethea (Saturniidae) and Malacosoma ameri- 
cana (Lasiocampidae) (see Brown, 1984). It has 
also been recorded as a hyperparasitoid of Apan- 
teles melanoscelus (Braconidae) and Anastatus 
disparts (Euplemidae) parasitizing larvae and 
eggs respectively of Lymantria dispar (Muese- 



beck & Dohanian, 1927; Howard & Fiske, 1911). 
Records of kuvanae as parasitoids of coreid eggs 
in Nigeria by Matteson (1981) are erroneous, 
being misidentifications of Ooencyrtus afer (see 
Prinsloo, 1987). 

Distribution. Japan, P.R. China, Korea, Tai- 
wan and also introduced into North America, 
Europe and north Africa. 

Material examined. Extralimital material. 
USA: 99, 110", USDA-EPL, from eggs of L. 
dispar, i.1977; 1$, Id", Gip(sy) Moth Labora- 
tory), No 4139B; CROATIA: 10$, 8cf, Vink- 
ovici, ex eggs Lymantria dispar, viii.1972-x.1974 
(J. Spaic); ALGERIA: 7$, Ain Mokra nr Bone, 
ex ova of Liparis dispar, em viii and x.1931, 
Ooencyrtus kuwanae How. Ch.Ferriere det.; 3$, 
Ain Mokra, ex Lymantria dispar, 1931 (J. de 
Lepiney); 3$, 2d", Chene-Liege Forest, ex eggs 
Lymantria dispar, ix.1984 (S. Hamza-Kroua); 
MOROCCO: 2$, 2d\ Mamora Forest, ex 
Lymantria dispar, 1931 (J. de Lepiney); JAPAN: 
19. Kyushu, Fukuoka, Hakozaki, ex egg Mala- 
cosoma neustria tartacea Motsch., 26. ii. 1959 
emerged 28. v. 1959 (Y. Miyatake). P.R. CHINA: 
13$, 3d", Beijing, ex eggs of lymantriid on 
poplar, x.1979; 5$, Heilongjiang, Heishan, ex 
eggs of Lymantria dispar L., 30.vii. 1976; Liaon- 
ing, Shenyang, ex eggs of Lymantria dispar L., 
17. ii. 1978 (Xiu Gongtian); 2$, Shanxi, 
Yuncheng, ex eggs of Stilpnotia salicis (L.), 
10. ix. 1975; 4$, Hebei, Funing, ex eggs of Den- 
drolimus spectabilis Butler, viii. 1987 (D.-x. 
Liao). Material in BMNH, IZAS. 

Comments. We have not seen any Oriental 
material of this species, but it is possibly the 
species recorded by Li et al. (1981) from the eggs 
of Lymantria xylina Swinhoe in Fujian Province 
(P.R. China) and it has been recorded from 
Taiwan (Trjapitzin, 1989). 

The species is most similar to O. phoebi (see 
comments under phoebi). 

Ooencyrtus dis sp.n. 

(Figs 54-58) 

Diagnosis. Female: face and scutellum bright 
metallic green; ocellar area and remainder of 
body blackish with a relatively weak metallic 
lustre; coxae blackish, femora, at least proximal 
two-thirds, especially dorsally, dark brown; 
remainder of legs yellowish; mandibles with 
three acute teeth (similar to Fig. 268); antennae 
(Fig. 54) with Fl-3 somewhat transverse and 
F4-6 longer than broad, clava about as long as 
F3-6 together, not conspicuously broader than 



28 



D. -W.HUANG AND J.S. NOYES 



funicle segments, with sutures parallel and with 
sensory area at extreme apex only; frontovertex 
one-fifth head width; scutellum with uniformly 
fine, punctate reticulate sculpture, only its 
extreme vertical apex smooth and shiny; oviposi- 
tor slightly exserted, exserted part about one- 
fifth as long as gaster. 

Female. Length, excluding ovipositor, about 
1.1 mm (holotype 1.13 mm). 

Head with ocellar area blackish but with weak 
metallic purple lustre, anterior to this distinctly 
metallic green, face bright metallic green, inter- 
antennal prominence purple, genae brassy- 
purplish; antennae with radicle, pedicel brown, 
scape testaceous-yellow, flagellum testaceous; 
pronotum, mesoscutum and axillae blackish with 
a moderate metallic green sheen; scutellum com- 
pletely bright metallic green; mesopleuron black; 
propodeum blackish; legs with coxae and proxi- 
mal two-thirds of femora, especially dorsally, 
dark brown; rest of legs yellow; wings hyaline, 
venation testaceous yellow; gaster black with 
metallic blue-green sheen on basal tergite; ovi- 
positor sheaths brown. 

Head in ocellar area with fine, raised reticulate 
sculpture, genae with longitudinally elongate 
sculpture; ocelli forming an angle of about 45°; 
the posterior ocelli separated from occipital mar- 
gin by their own lengths and nearly touching eye 
margins; inner eye orbits strongly divergent ven- 
trally (Fig. 56); frontovertex one-fifth head 
width; eyes with conspicuous short setae; occipi- 
tal margin hardly rounded; head in facial view 
nearly as broad as high; antennae inserted just 
below ventral eye margins (Fig. 56), toruli sepa- 
rated from mouth margin by less than their own 
lengths; antennae (Fig. 54) with pedicel plus 
flagellum together about one-fifth longer than 
head width; pedicel nearly as long as Fl-3 
together which are somewhat transverse, F4-6 
longer than broad; clava about as long as F3-F6 
together, not conspicuously broader than funicle 
segments, its sutures parallel and with sensory 
area limited to extreme apex only; mandibles 
with three sharp teeth (similar to Fig. 268). 
Relative measurements (holotype): head width 
42, head height 40, minimum frontovertex width 
8, POL 3, OPL 3.5, OOL 0.1, eye length 29, eye 
width 25, malar space 13, scape length 20, scape 
width 4.5; other proportions of the antennae as 
in Fig. 54. 

Mesoscutum with shallow imbricate sculpture; 
scutellum with more or less uniform, fine, punc- 
tate reticulate sculpture, only its vertical apex 
smooth and shiny; fore wing with venation and 
distribution of setae basally as in Fig. 55. Rela- 



tive measurements (holotype): forewing length 
93, forewing width 42; hindwing length 67; hind- 
wing width 17. 

Gaster, excluding ovipositor, distinctly shorter 
than thorax; ovipositor (Fig. 58) exserted, 
exserted part about one-fifth as long as gaster, 
gonostylus about one-fifth ovipositor length; 
hypopygium as in Fig. 57. Relative measure- 
ments: ovipositor length 48, gonostylus 11 [mid 
tibia 34]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
9, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, ii.1985, FIT (J.S. Noyes). 
Paratype, 1$, same data as holotype (BMNH). 

Comments. Females of Ooencyrtus dis can be 
separated from other species of the genus by the 
combination of the tridentate mandibles, 
strongly divergent inner orbits, totally green 
scutellum which is uniformly finely punctate 
reticulate and the dark brown coxae and femora. 

Ooencyrtus hercle sp.n. (Figs 58-63) 

Diagnosis. Body including gaster dark brown 
or black with a weak metallic blue sheen on 
mesoscutum, apex of scutellum more strongly 
shining blue or purple; legs with all coxae, 
femora except base and apex brown; tibiae 
mostly testaceous; mandible with one tooth and a 
broad truncation; scutellum with deeper sculp- 
ture than meoscutum; forewing (Fig. 60) with 
relatively dense setae in basal cell and dense fine 
setae on ventral side near hind margin; postmar- 
ginal vein as long as stigmal vein. Female: anten- 
nae (Fig. 59) with Fl-3 subequal and 
subquadrate, conspicuously smaller than follow- 
ing ones, F6 somewhat transverse; clava as long 
as F3-6 together, sensory area limited to apex 
only; frontovertex one-third head width; oviposi- 
tor hidden or very slightly exserted. Male: anten- 
nae with all funicle segments slightly longer than 
broad and with setae a little longer than diameter 
of segments; frontovertex about two-thirds head 
width and aedeagus about two-thirds as long as 
mid tibia. 

Female. Length 0.5-0.6 mm, (holotype 0.57 
mm). 

Head dark brown or blackish with a weak 
metallic blue tinge; antennae with radicle, scape 
and pedicel blackish, flagellum brown testa- 
ceous; thorax dark brown or blackish with a 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



29 



metallic blue sheen, scutellum anteriorly cop- 
pery, posterior quarter or so blue or purple; 
mesopleuron dark brown; legs with coxae and 
femora dark brown, base and apex of femora 
testaceous, tibia with basal two-thirds dark 
brown, distal one-third testaceous; wings hya- 
line, venation testaceous; gaster dark brown 
blackish with a slight metallic blue sheen; ovi- 
positor sheaths brown. 

Frontovertex with fine, raised reticulate sculp- 
ture; sculpture on genae longitudinally elongate; 
ocelli more or less forming a right triangle; 
posterior ocelli separated from occipital margin 
by their own lengths and distinctly separated 
from eye margins; frontovertex about one-third 
head width as in Fig. 62 (holotype a little more 
than one-third); eyes not conspicuously hairy; 
occipital margin rounded; toruli separated from 
mouth margin by slightly less than their own 
lengths; antennae (Fig. 59) with pedicel plus 
flagellum about as long as head width; pedicel 
slightly longer than Fl-2 together; funicle seg- 
ments slightly enlarged towards apex; Fl-3 sub- 
equal and subquadrate, conspicuously smaller 
than F4-6; clava as long as F3-6 together, its 
sutures parallel and apex rounded with sensory 
area limited to extreme apex only; mandible with 
one tooth and a broad truncation. Relative mea- 
surements (holotype): head width 25, head 
height 23, minimum frontovertex width 9, POL 
3, OPL 1.5, OOL 0.7, eye length 15, eye width 
12, malar space 8.5, scape length 11.5, scape 
width 2, other proportions of the antennae as in 
Fig. 59. 

Mesoscutum with shallow imbricate sculpture; 
scutellum with conspicuously deeper reticulate, 
polygonal sculpture which is more elongate later- 
ally, the extreme posterior margin smooth and 
shiny; fore wing venation and distribution of 
setae basally as in Fig. 60; basal cell with dense 
setae on its ventral side near posterior wing 
margin; postmarginal vein as long as stigmal 
vein. Relative measurements (holotype): forew- 
ing length 65, forewing width 29; hindwing length 
37; hindwing width 10. 

Gaster shorter than thorax; ovipositor (similar 
to Fig. 86) with second valvifers basally moder- 
ately curved with curved part relatively long; 
gonostylus about one-fifth ovipositor length; 
hypopygium as in Fig. 61. Relative measure- 
ments: ovipositor length 83, gonostylus 18 [mid 
tibia 60]. 

Male. (Length 0.54 mm). Generally similar to 
female but for antennae, relatively wider fron- 
tovertex and genitalia. Antennae (Fig. 63) with 
all funicle segments longer than broad, pedicel 



and flagellum together distinctly longer than 
head width; longest setae slightly longer than 
diameter of segments; frontovertex two-fifths 
head width; aedeagus about two-thirds as long as 
mid tibia, digiti moderately elongate and each 
with a single apical hook. 

Hosts. Recorded below as a parasitoid of the 
eggs of Rondotia menciana (Lepidoptera: Bom- 
bycidae). 

Distribution. P. R China. 

Material examined. Type material. Holotype 
9, PR. CHINA: Zejiang, Wuxing, ex eggs of 
Rondotia menciana Moor, v. 1956 (Fuqi Zhang) 
(IZAS). Paratypes, 9$, 3d\ same data as holo- 
type (IZAS, BMNH). 

Non type material. A further 40 unmounted 
specimens in poor condition, but with same data 
as type material (IZAS). 

Comments. Ooencyrtus hercle is close to cor- 
betti and can be separated from this species by its 
smaller size, relative proportions of funicle seg- 
ments and wider frontovertex in the female. 

Ooencyrtus corbetti Ferriere 

(Figs 64-66) 

Ooencyrtus corbetti Ferriere, 1931: 284. LECTO- 
TYPE <J> (here designated), Malaysia (BMNH. 
examined). 

Diagnosis. Body including gaster black, weakly 
metallic; interantennal prominence purple; 
scutellum coppery in basal half, its apical half 
weakly to strongly metallic green; legs with all 
coxae, femora except both base and apex black; 
tibiae normally totally yellow but occasionally 
fore tibia with a faint brown subbasal ring; scutel- 
lum (similar to Fig. 80) with fairly regular, 
slightly elongate, reticulate sculpture which is 
conspicuously deeper than that on mesoscutum, 
its extreme posterior margin smooth and shiny; 
mandible (similar to Fig. 69) with two acute teeth 
and a truncation; forewing (Fig. 65) with post- 
marginal vein a little shorter than stigmal vein. 
Female (0.80-1.93 mm): antennae (Fig. 64) with 
scape slightly expanded in the middle, about 4 
times as long as broad; pedicel as long as Fl-2; 
Fl-2 subequal and hardly longer than broad, 
conspicuously smaller than F3-6 which are dis- 
tinctly longer than broad; clava as long as F4-6 
together or a little longer, its apex rounded and 
with sensory part limited to extreme apex only; 
frontovertex about one-quarter head width; ovi- 
positor hidden, the sheaths brown; basal part of 
second valvifers relatively broad (Fig. 66). Male 



30 



D. -W.HUANG AND J.S. NOYES 



(length about 0.8 mm): antennae with all funicle 
segments longer than broad, pedicel and flagel- 
lum together distinctly longer than head width; 
longest setae about twice as long as diameter of 
segments; frontovertex slightly more than two- 
fifths head width. 

HOSTS. Reared from eggs of Podontia quatuor- 
decimpunctata (Coleoptera: Chrysomelidae). 

Distribution. Malaysia. 

Material examined. Type material. Lecto- 
type $, MALAYSIA: Kuala Lumpur, from eggs 
of Podontia 14-punctata, 5. VII. 1921, (G. H. Cor- 
bett), B.M. TYPE HYM. 5. 1,072, also with a 
red 'type' label. Paralectotypes, 7$, each with a 
yellow 'Cotype' label, otherwise same data same 
as lectotype; 1$, with a yellow 'Paratype' label, 
otherwise as lectotype (BMNH). 

Comments. Ooencyrtus corbetti is very close to 
podontiae and may eventually prove to be syn- 
onymous. However we have seen only limited 
material and for the present we prefer to retain 
the two as distinct because of small differences in 
setation at the base of the forewing (cf. Figs 65 
and 70) and in the relative proportions of the 
antennal segments. In females of corbetti only 
Fl-2 are relatively small and the clava as long as 
F4-6 combined, whilst in podontiae Fl-3 are 
subequal and relatively small and the clava is as 
long as F3-6 combined. 

Ooencyrtus podontiae (Gahan) 
(Figs 67-73) 

Schedius podontiae Gahan, 1922: 51, $. LEC- 
TOTYPE $ (here designated), Indonesia: 
Java (USNM, examined). 

Ooencyrtus podontiae (Gahan); Ferriere, 1931: 
282, 286. 

Diagnosis. Body including gaster black, weakly 
metallic; interantennal prominence purple; 
scutellum coppery in basal half, its apical half 
weakly to strongly metallic green; legs with all 
coxae, femora except both base and apex black; 
tibiae yellow; scutellum (Fig. 71) with fairly 
regular, slightly elongate, reticulate sculpture 
which is conspicuously deeper than that on 
mesoscutum, its extreme posterior margin 
smooth and shiny; mandible (Fig. 69) with two 
acute teeth and a truncation; forewing (Figs 70) 
with postmarginal vein shorter than stigmal vein. 
Female (0.80-1.00 mm): antennae (Figs 67, 68) 
with scape slightly expanded in the middle, 2>-A 
times as long as broad; pedicel as long as Fl-2; 
Fl-3 subequal and quadrate, F4-F6 longer than 



broad, subequal; clava as long as F3-6, its apex 
rounded and with sensory part limited to extreme 
apex only; frontovertex about one-quarter head 
width; ovipositor hidden to slightly exserted, the 
sheaths brown; basal part of second valvifers 
relatively broad (Fig. 72); hypopygium as in Fig. 
73. Male (length about 0.8 mm): antennae with 
all funicle segments longer than broad, pedicel 
and flagellum together distinctly longer than 
head width; longest setae about twice as long as 
diameter of segments; frontovertex about two- 
fifths head width. 

Hosts. Reared from eggs of Podontia quatuor- 
decimpunctata (Coleoptera: Chrysomelidae). 

Distribution. Indonesia. 

Material examined. Type material. Lecto- 
type 9, INDONESIA: Java, Buitenzorg, 
27.XI.1919 (S. Leefmans) (USNM). Paralecto- 
types of podontiae: 7§, same data same data as 
the lectotype (USNM, BMNH). The type series 
bears a label giving the eggs of Podontia 
14-punctata (= quatuordecimpunctata) as the 
host, even though Gahan originally gives the host 
as Podontia affinis Grond. 

Other material. INDONESIA: 1$, Java, from 
same series as lectotype of Schedius podontiae; 
1§, Sulawesi Utara, Dumoga-Bone NP, Toraut, 
200m, BMNH Canopy Fog #15, 19.vii.1985; 1$, 
BMNH Canopy Fog #13, ll.vii.1985. Material 
in USNM, BMNH 

Comments. Ooencyrtus podontiae is close to 
corbetti, plautus and pindarus (see comments 
under those species). 

Ooencyrtus pindarus sp.n. 

(Figs 74-82) 

Diagnosis. Mandible (Fig. 75) with two teeth 
and a truncation; postmarginal vein a little 
shorter than stigmal vein. Female: body includ- 
ing gaster black with a weak bluish lustre on the 
head and mesoscutum, this stronger on face, 
interantennal prominence purple; scutellum cop- 
pery basally, apically metallic green to purple at 
extreme apex; legs with all coxae and femora 
blackish; tibiae yellow but partly dark brown; 
head triangular in side view; antennae (Fig. 76) 
with scape subcylindrical, Fl-2 subequal and 
transverse, conspicuously smaller than F3-6; 
clava slightly longer than F4-6 together, sutures 
parallel, apex rounded and sensory area limited 
to extreme apex only; frontovertex one-quarter 
head width; eyes as long as broad, or broader; 
scutellum (Fig. 80) with relatively deep longitudi- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



31 



nally reticulate sculpture, only its extreme poste- 
rior margin smooth; ovipositor hidden or very 
slightly exserted. Male: generally similar to 
female, but body much duller, dark purplish 
brown or blackish with hardly a metallic sheen, 
legs slightly darker; antenna (Fig. 82) with all 
funicle segments slightly longer than broad and 
clothed in setae, the longest of which are a little 
longer than diameter of segments. 

Female. Length 0.65-0.85 mm, (holotype 0.76 
mm). 

Head in ocellar area blackish, faintly purple, 
anterior to this weak metallic blue, between 
scrobes and eyes blue or green, interantennal 
prominence purple; antennae with radicle, scape 
and pedicel dark brown, flagellum testaceous; 
pronotum, mesoscutum and axillae blackish with 
a slight metallic blue tinge; scutellum dark cop- 
pery purple in basal half or so, apex and sides 
metallic green, extreme apex purple; mesopleu- 
ron black; propodeum blackish; legs with coxae 
and femora black; apices of femora yellowish; 
fore tibia yellow, medially dark brown; mid and 
hind tibia each with a subbasal dark brown ring; 
wings hyaline, venation testaceous; gaster black 
with metallic blue sheen; ovipositor sheaths dark 
brown. 

Head in ocellar area with fine, raised reticulate 
sculpture; sculpture on genae longitudinally 
elongate; ocelli forming about a 45° angle; poste- 
rior ocelli separated from occipital margin by 
their own lengths and slightly separated from eye 
margins; frontovertex one-quarter head width; 
eyes not conspicuously hairy, very large, as long 
as broad or broader; occipital margin rounded; 
head in front view (Fig. 74) slightly broader than 
long, almost triangular in side view, quite similar 
in shape to species of Adelencyrtus and related 
genera; torulus separated from mouth margin by 
less than its own length; antennae (Fig. 76) with 
pedicel plus flagellum about as long as head 
width; pedicel slightly longer than Fl-2 together 
which are subequal and transverse and conspicu- 
ously smaller than F3-6; clava slightly longer 
than F4-6 together, sutures parallel and apex 
rounded with sensory area restricted to extreme 
apex only; mandible (Fig. 75) with two teeth and 
a truncation. Relative measurements (holotype): 
head width 30, head height 27, minimum fron- 
tovertex width 7.5, POL 3, OPL 2.5, OOL 0.5, 
eye length 18, eye width 17, malar space 10, 
scape length 11, scape width 2.5; other propor- 
tions of the antennae as in Fig. 76. 

Mesoscutum with shallow imbricate sculpture; 
scutellum (Fig. 80) with deeper, more or less 
uniform, longitudinally reticulate sculpture, only 



its posterior margin smooth, its apex pointed; 
fore wing with venation and distribution of setae 
basally as in Fig. 77. Relative measurements 
(holotype): forewing length 70, forewing width 
28; hindwing length 48; hindwing width 10. 

Gaster acute apically with ovipositor (Fig. 78) 
hidden or very slightly exserted, hypopygium as 
in Fig. 81. Relative measurements (paratype): 
ovipositor length 98, gonostylus 20 [mid tibia 69]. 

Male. Similar to female but differs as follows. 
Body generally dark purplish brown or blackish, 
hardly metallic; legs slightly darker; head in side 
view more or less evenly rounded anteriorly; 
frontovertex one-third head width; antennae 
(Fig. 82) with all funicle segments longer than 
broad and clothed in relatively long setae; genita- 
lia (Fig. 79) with aedeagus about half as long as 
mid tibia. 

Variation. Very little in material available, but 
in females the tibiae may be largely dark brown 
in basal half or only with a narrow dark brown 
ring; in the males the tibiae may be more or less 
completely dark brown or testaceous-yellow in 
apical half. 

Hosts. Reared from eggs of Plesispa reichei 
(Coleoptera: Chrysomelidae). 

Distribution. Malaysia, Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Java, Selangor (? - see com- 
ments below), Mojoagung, from eggs of Plesispa 
reichei, 19.vii.1989, CIE 20619 (Wornoto) 
(BMNH). Paratypes, MALAYSIA: 30$, same 
data as holotype; 39$, 6cf , Penang I. to Melaka, 
ex eggs of Plesispa reichei, 13-27. xii. 1990 (R. & 
L. Burkhart); 159, 6o\ Sumatra, ex eggs of 
Plesispa reichei, 11.1936 (R. Aurbowo); 7$, 
Sulawesi, ex eggs of P. reichei, 1989, CIE 
A20420; 2$, Sulawesi, Manado, ex. eggs of P. 
reichei, 30.x. 1989, CIE A20715. Material in 
BMNH, BPBM, IZAS, PPRI, USNM, ZISP, 
QMB. 

Comments. We are unable to localise the series 
from which the holotype is designated. The label 
on the specimens reads both Java and Selangor 
which cannot be correct since Selangor is part of 
Malaysia and not Indonesia. International Insti- 
tute of Entomology (= CIE) records specify that 
the material was originally collected in Indone- 
sia. However we are unable to find anywhere 
called Mojoagung in Indonesia, although there is 
an island called Mojo near Sumbawa to the east 
of Java. 

Ooencyrtus pindarus is very close to corbetti 



32 



D. -W.HUANG AND J.S. NOYES 



and podontiae but can be separated by the head 
of the female being distinctly triangular in side 
view, the broad eyes and the tibiae being largely 
dark brown. In the other species the head is more 
or less evenly curved anteriorly in side view, the 
eyes are clearly longer than broad and the tibiae 
are entirely yellow or nearly so. The head shape 
in pindarus is very probably a function of the 
shape of its host, the egg of P. reichei being 
flattened and elongate. This would produce adult 
parasitoids with a depressed body, triangular 
head and relatively broad eyes. This suggests that 
the above material may be only a host-induced 
form of podontiae. However, the consistent dif- 
ferences in leg coloration and overlapping ranges 
lead us to believe that the material must belong 
to a distinct species. 

Ooencyrtus plautus sp.n. 

(Figs 83-88) 

Diagnosis. Body including gaster black, weakly 
metallic blue on head and mesoscutum; legs with 
coxae and femora brown; tibiae yellow but with a 
distinct subbasal brown ring; antennae (Figs 83, 
88) of both sexes with scape slightly expanded in 
middle; scutellum (similar to Fig. 80) with rela- 
tively deep elongate-reticulate sculpture only the 
extreme posterior margin smooth; mandible 
(similar to Fig. 75) with two teeth and a trunca- 
tion. Female: antenna with Fl-3 subequal and 
subquadrate, conspicuously smaller than F4-6; 
clava about as long as F3-6 together; frontover- 
tex about one-quarter head width; ovipositor 
hidden or hardly exserted. Male: frontovertex 
about one-third head width; all funicle segments 
at least about 2 times as long as broad and with 
longest setae about 4 times the diameter of any 
segment, otherwise similar to female. 

Female. Length 0.65-0.8 mm (holotype 0.70 
mm). 

Head blackish, weakly metallic in ocellar area, 
face and genae weakly metallic green, interan- 
tennal prominence purplish; antennae with 
radicle, scape except distal end and pedicel dark 
brown or blackish, flagellum testaceous; prono- 
tum and axillae blackish with a very slight purple 
tinge; mesoscutum similar but bluish; scutellum 
basally black with purple-brown sheen, its apex 
and sides narrowly metallic green, the extreme 
apex purple; mesopleuron dark brown tinged 
purple; legs with coxae and femora dark brown; 
apices of femora yellowish; tibiae yellow, with a 
subbasal brown ring; wings hyaline, venation 
testaceous; gaster black with slight bluish sheen; 
ovipositor sheaths dark brown. 



Head in lateral view not triangular and more 
or less evenly curved anteriorly; ocellar area with 
fine, raised reticulate sculpture; genae relatively 
shiny and with longitudinally elongate sculture; 
ocelli forming a right angle, posterior ocelli sepa- 
rated from occipital margin by less than their 
own length and slightly separated from eye mar- 
gins; frontovertex slightly more than one-quarter 
head width; eyes not conspicuously hairy, clearly 
longer than broad; occipital margin hardly 
rounded; toruli separated from mouth margin by 
less than their own lengths (Fig. 85); antennae 
(Fig. 83) with pedicel plus flagellum about as 
long as head width; clava with apex rounded, its 
sutures parallel and sensory area at extreme apex 
only; mandible with two teeth and a truncation. 
Relative measurements (holotype): head width 
32, head height 30, minimum frontovertex width 
9, POL 4.5, OPL 1.5, OOL 0.3, eye length 21, 
eye width 18, malar space 10, scape length 12, 
scape width 3; other proportions of the antennae 
as in Fig. 83. 

Mesoscutum with shallow imbricate sculpture; 
scutellum (similar to Fig. 80) with deeper, longi- 
tudinally reticulate sculpture, only its extreme 
lateral and posterior margins smooth and shiny; 
forewing with venation and distribution of setae 
basally as in Fig. 84. Relative measurements 
(holotype): forewing length 68, forewing width 
31; hindwing length 50; hindwing width 12. 

Gaster with ovipositor (Fig. 86) hidden or 
hardly exserted, basal part of second valvifer 
relatively narrow and strongly curved, gonostylus 
about one-fifth ovipositor length; hypopygium 
similar to Fig. 81. Relative measurements 
(paratype): ovipositor length 107, gonostylus 24 
[mid tibia 80]. 

Male. Length 0.6-0.65 mm. 

Very similar to female differing slightly in leg 
coloration, relative width of the frontovertex and 
forewing and structure of antennae and genitalia. 
Tibiae with subbasal brown ring relatively broad, 
on mid tibia nearly two thirds its length; fron- 
tovertex at least two-fifths head width; antennae 
(Fig. 88) with pedicel conspicuously shorter than 
first funicle segment, each funicle segment with 
very long setae around the middle, other propor- 
tions of antennae as in Fig. 88; forewing broader, 
about 1.9 times as long as broad; genitalia as in 
Fig. 87 with aedeagus a little more than half 
length of mid tibia. 

Variation. Very little in material available. 
The frontovertex of the male varies in relative 
width from about two-fifths head width in larger 
specimens to about one-half in smaller ones. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



33 



Hosts. Recorded below from eggs of Papilio 
citri (lapsus for P. demoleus Linnaeus?) and 
Papilio sp. (Lepidoptera: Papilionidae). 

Distribution. India. 

Material examined. Type material. Holotype 
9, INDIA: Karnataka, Bangalore, ex. eggs of 
Papilio citri, CIE A19148, viii.1986 (BMNH). 
Paratypes, INDIA: 4$, 3cf, Karnataka, Banga- 
lore, ex Papilio sp., 13.vii.1987, CIE 19148. 
BMNH, IZAS. 

Comments. Females of Ooencyrtus plautus are 
similar in general appearance to those of podon- 
tiae and pindarus, all three species having the 
scape conspicuously expanded medially, similar 
antennal structure and sculpture on the scutel- 
lum. O. plautus can be separated from podontiae 
by the presence of distinct brown rings on the 
tibiae, whilst it can be distinguished from pin- 
darus by the head shape in combination with the 
forewing venation. The head of pindarus is dis- 
tinctly triangular in side view and the stigmal vein 
of plautus is relatively longer than that of pin- 
darus (cf. Figs 77 and 84). On the other hand 
males of these species can be separated easily on 
the relative length of the setae on the flagellum 
and the shape of the funicle segments (compare 
Figs 82 and 88). 

Ooencyrtus daphne sp.n. 

(Figs 89-93) 

Diagnosis. Female: antennae with distal two- 
thirds of scape and distal three funicle segments 
whitish other segments brown; head and thorax 
blackish, but head with a slight metallic green 
sheen, frontovertex purplish, mesoscutum and 
scutellum with a distinct purplish lustre; all coxae 
and fore and mid femora dark brown; hind 
femora totally yellow; fore-tibia partly brown; 
mid and hind tibia yellow; tarsi testaceous; gaster 
basally both dorsally and ventrally yellow, later- 
ally and apically dark brown; antennae (Fig. 89) 
with Fl-3 subequal and subquadrate and smaller 
than F4-6 which are larger and a little longer 
than broad; scutellum entirely smooth except for 
a small triangular basal area; forewing with a 
median cloudy band (Fig. 90); ovipositor charac- 
teristic (Fig. 93), very broad and much shorter 
than mid-tibia, hypopygium produced apically in 
the form of a short tongue, or mucro (Fig. 92), 
which reaches apex of gaster. Male: unknown. 

Female. Length about 0.8-0.9 mm (holotype 
0.8 mm). 

Head black, in ocellar area and interantennal 



prominence with a metallic purplish tinge; occipi- 
tal, scrobal area and genae with a distinct metal- 
lic blue-green tinge; antennae with radicle and 
proximal one-third of scape blackish, distal two- 
third of scape pale yellow, pedicel and Fl-2 or F3 
brown, F4-6 whitish, clava brown; dorsum of 
thorax purplish reflection, scutellum a little 
brighter than mesoscutum; mesopleuron and 
propodeum dark brown; coxae concolorous with 
mesopleuron; fore and midfemora dark brown; 
hind femora totally yellow; fore tibia proximally 
brown, distal half yellow; mid and hind tibia 
yellow; tarsi testaceous-orange; forewing with a 
cloudy band across middle (Fig. 90); gaster ante- 
riorly yellow both dorsally and ventrally, later- 
ally and apically dark brown; ovipositor sheaths 
dark brown. 

Head with fine, regularly reticulate sculpture 
in ocellar area, sculpture on genae longitudinally 
elongate; ocelli forming a slightly acute angle, 
the posterior ocelli separated from occipital mar- 
gin by less than their own lengths and conspicu- 
ously separated from eye margins; frontovertex 
about one-quarter head width; eyes fairly con- 
spicuously hairy; antennae inserted just below 
the ventral eye margin, toruli separated from 
mouth margin by less than their own lengths; 
antennae (Fig. 89) with pedicel plus flagellum 
about as long as head width; pedicel a little 
longer than following two segments together; 
F1-F3 slightly transverse, F5-F6 at least slightly 
longer than broad, subquadrate; clava about as 
long as F2-6 together, the outer suture hardly 
oblique and the apex with a very short oblique 
apical truncation, the sensory area relatively 
small and limited to apex only; mandibles (Fig. 
91) with three teeth, but the third, upper tooth 
somewhat truncate. Relative measurements 
(holotype): head width 38, head height 36, mini- 
mum frontovertex width 10, POL 6, OPL 2, 
OOL 1, eye length 26 eye width 20, malar space 
10, scape length 16, scape width 3; other propor- 
tions of the antennae similar to Fig. 89. 

Mesoscutum with shallow imbricate sculpture; 
scutellum entirely smooth but for a small rugu- 
lose trianglar area at base, its apex broadly 
rounded; forewing with venation and distribution 
of setae basally as in Fig. 90, costal cell relatively 
narrow and linea calva open posteriorly. Relative 
measurements (holotype): forewing length 86, 
forewing width 36, hindwing length 60, hindwing 
width 14. 

Gaster with ovipositor hidden or slightly 
exserted; ovipositor (Fig. 93) very broad, robust, 
gonostylus robust, sharply tapering towards tip 
and about one-quarter as long as ovipositor; 
hypopygium (Fig. 92) with a short, median, 



34 



D. -W.HUANG AND J.S. NOYES 



apical projection which reaches the apex of the 
gaster. Relative measurements (paratype): ovi- 
positor length 58, gonostylus 15 [mid tibia 79]. 

Male. Unknown. 

Variation. There is some variation in the col- 
oration of the antennal segments. In the holotype 
only Fl-2 are brown, whereas in the paratypes 
Fl-3 are distinctly brown. 

Hosts. Unknown. 

Distribution. Brunei, Malaysia. 

Material examined. Type material. Holotype 
$, BRUNEI: Ulu Temburong, 300m. ii-iii.1982 
(M. Day) (BMNH). Paratypes, BRUNEI: 1$, 
Bukit Sulang nr. Lamunin, BM 1982-388, 
20.viii-10.ix.1982 (N.E. Stork); MALAYSIA: 
1$, Mt Kinabalu NP, Poring Hot Springs, 490m, 
BM 1991-85, 20.viii.1988 (A. Smetana). Both 
paratypes in BMNH. 

Comments. Ooencyrtus daphne and flavipes 
belong to the same species group which was 
given separate generic status by Timberlake as 
Xesmatia. The group is characterised by triden- 
tate mandibles with upper tooth rounded or 
truncate, smooth shiny scutellum, relatively 
robust ovipositor and extended hypopygium 
which reaches apex of gaster. The structure of 
the ovipositor and hypopygium are unique within 
Ooencyrtus but we do not believe that these 
differences require the recognition of Xesmatia 
as a distinct genus because it is highly likely that 
these two species represent only a derived lin- 
eage within Ooencyrtus. 

Ooencyrtus daphne can be separated from fla- 
vipes by having all coxae dark brown and a 
cloudy band across the forewing. In flavipes at 
least the hind coxae are yellow, the wings are 
completely hyaline and the marginal- 
postmarginal vein has only two long setae. 

Ooencyrtus flavipes (Timberlake) 
(Figs 94-102) 

Xesmatia flavipes Timberlake, 1920: 425-128, $. 

Holotype 9, Hawaiian Is. (BPBM, examined). 
Ooencyrtus flavipes (Timberlake); Noyes & 

Hayat, 1984: 309. 

Diagnosis. Female (length: 0.48-1.03 mm): 
antennae with scape yellow to testaceous, flagel- 
lum normally bicolorous, clava brown and 
funicle segments testaceous, occasionally unicol- 
orous brown; head and thorax blackish, but head 
with a slight metallic green sheen, frontovertex 
purplish, mesoscutum and scutellum with a dis- 



tinct purplish or bluish lustre; fore coxae dark 
brown, mid coxae variable, hind coxae yellow; 
gaster normally mostly yellow with apex and 
sides dark brown; scutellum entirely smooth 
except for a small triangular, rugulose area at 
base; mandibles with three obtuse teeth; oviposi- 
tor robust (Fig. 100); hypopygium (Figs 101, 102) 
with a short median posterior projection which 
reaches apex of gaster; gaster mainly yellowish 
with dorsolateral margin brown. 

Variation. There is some notable variation in 
coloration of the gaster, relative width of the 
frontovertex and proportions of the antennal 
segments. The majority of specimens have the 
gaster mostly yellow laterally and apically dark 
brown. The brown coloration may be reduced to 
a narrow line or more extensive, the gaster 
almost entirely dark brown. Larger specimens 
have the frontovertex relatively wider (0.4 times 
head width in larger specimens to 0.25 times 
head width in smaller ones) and longer funicle 
segments (compare Figs 96 and 97). There is also 
some variation in strength of the sculpture on the 
scutellum, usually most of the scutellum is com- 
pletely smooth and shiny but some specimens are 
largely shallowly reticulate. The shape of the 
hypopygium varies from relatively transverse 
(Fig. 102) to relatively long (Fig. 101). We think 
that this is probably due to differences in the 
degree of sclerotisation of the membranous part 
connecting the hypopygium to the fourth gastral 
sternite. 

Hosts. Unknown. 

Distribution. Nepal, Thailand, Brunei, Indo- 
nesia, Hawaii. 

Material examined. Type material. Holotype 
9, HAWAII: Palolo Crater, Oahu, 20.X.1918 
(J.C. Bridwell, nee 20.xii.1918), with one forew- 
ing, one hindwing, one antenna and one man- 
dible on slide (BPBM). 

Other material. NEPAL: 1$, Kathmandu, 
1350m, vi. 1984 (M.G. Allen); THAILAND: 1$, 
Nakhon Rat Chasima, Khao Yai NP 700-800m, 
11-18. iv. 1990 (E. Fuller); BRUNEI: 1$, Ulu 
Temburong, 300m, ii-iii.1982 (M. Day); 1$, 
Bukit Sulang, Nr. Lamunin, 20. viii— lO.ix. 1982 
(N.E. Stork); INDONESIA: 1$, Sulawesi 
Utara, Dumoga-Bone NP, Toraut, 8.ii.l985 
(BMNH Canopy Fog #3); 7$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, iv.1985 (J.S. Noyes); 
6$, Sulawesi Utara, Dumoga-Bone NP, Toraut, 
v.1985 (J.H. Martin); 1$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, v. 1985 (A. D. Aus- 
tin); 4$, Sulawesi Utara, Dumoga-Bone NP 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



35 



Gunung Mogoganipa, 1000m, v. 1985 (J.S. Noyes 
& J.H. Martin); 19- Sulawesi Utara, Danau 
Mooat (Kotomobaqu), v. 1985 (J.S. Noyes); 4$>, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
500m, 16-23, vi.1985 (A.D. Austin); 1$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
vi.1985 (A.D. Austin); 4$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 200m, ll.vii.1985, 
(BMNH Canopy Fog #13); 1$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 19.vii.1985, (BMNH 
Canopy Fog #15); 1$, Sulawesi Utara, Dumoga- 
Bone NP, Gunung Mogogonipa, 1000m, xi.1985. 
Material in BMNH, IZAS, ZISP, USNM, PPRI, 
CNCO. 

Comments. Ooencyrtus flavipes is very close to 
daphne (see comments under O. daphne). 

Ooencyrtus dione sp.n. 

(Figs 103-105) 

Diagnosis. Female: head and scutellum dark 
brown with a distinct metallic green sheen, 
mesoscutum with a blue sheen, gaster purplish 
brown, rest of body dark brown; all coxae, fore 
femora and hind femora at least partly black, mid 
leg and all tibiae entirely yellow or nearly so; 
mandibles with one tooth and a very broad 
truncation; antennae dark brown with F5 and F6 
yellow; antennae (Fig. 104) with scape extremely 
enlarged at middle, about 2.5 times as long as 
broad; flagellum strongly clavate, Fl slightly 
longer than broad, F2 subquadrate, F3-6 con- 
spicuously transverse; clava conspicuously 
enlarged and much wider than F6, its sutures 
subparallel and sensory part enlarged but limited 
to apex only; head with frontovertex about one- 
quarter head width; scutellum with shallow sculp- 
ture, not deeper than that on mesoscutum or 
hardly so, posterior margin smooth and shiny 
(Fig. 105) and apex pointed; forewing with a 
conspicuous naked basal area (Fig. 103), with a 
complete row of setae on ventral side near poste- 
rior wing margin; stigmal vein long and slender, 
about twice as long as postmarginal vein; oviposi- 
tor hardly exserted, sheaths yellow. Male: 
unknown. 

Female. Length 1.37 mm (holotype). 

Head black, with a strong metallic green sheen 
in ocellar area; antennal scrobes dorsally purple, 
lower parts of scrobes and face green, interanten- 
nal prominence coppery purple; antennal radicle 
and scape black, flagellum black-brown with F5 
and F6 contrasting yellow; pronotum, mesoscu- 
tum and axillae black with a strong metallic blue 
sheen on mesoscutum; scutellum metallic green; 



tegulae, mesopleuron and propodeum dark 
brown; coxae dark brown; fore femora and 
proximal two third of hind femora dark brown; 
base and apical one third of hind femora yellow, 
mid legs yellow; fore tibiae with a faint dark 
brown subbasal ring; hind tibiae yellow; wings 
hyaline, venation testaceous-yellow; gaster dark 
purple-brown with a metallic blue sheen on basal 
tergites; ovipositor sheath yellow. 

Head in ocellar area with regular, raised 
reticulate sculpture; sculpture on post ocellar 
area transverse; lower face and genae more shiny 
than frontovertex, sculpture longitudinally elon- 
gate; ocelli forming a slightly acute angle; poste- 
rior ocelli separated from occipital margin by 
about their own lengths and distinctly separated 
from eye margins; frontovertex a little more than 
one-quarter head width; eyes inconspicuously 
hairy; occipital margin slightly rounded; anten- 
nae hardly inserted below ventral eye margins; 
toruli separated from mouth margin by less than 
their own lengths; antennae (Fig. 104) with 
pedicel plus flagellum 0.9 times head width; 
scape extremely enlarged at middle, about 2.5 
times as long as broad; flagellum strongly clav- 
ate; clava distinctly broader than F6, its sutures 
subparallel and sensory area at apex only; man- 
dibles with one tooth and a broad truncation 
(similar to Fig. 213). Relative measurements 
(holotype): head width 58, head height 51, mini- 
mum frontovertex width 17, POL 8.5, OPL 4, 
OOL 1.5, eye length 35, eye width 26, malar 
space 20, scape length 22, scape width 9; other 
proportions of antennae as in Fig. 104. 

Mesoscutum with shallow, fine, more or less 
polygonally reticulate or imbricate sculpture; 
sculpture on scutellum similar but gradually 
becoming shallower towards apex and sides so 
that apex is completely smooth and shiny and 
pointed (Fig. 21); forewing with venation and 
distribution of setae basally as in Fig. 103; stig- 
mal vein long and slender, about twice as long as 
postmarginal vein. Relative measurements (holo- 
type): forewing length 105, forewing width 46; 
hindwing length 72; hindwing width 18. 

Gaster shorter than thorax; ovipositor (similar 
to Fig. 24) hardly exserted, with gonostylus 
about one-sixth as long ovipositor; hypopygium 
similar to Fig. 25. Relative measurements 
(paratype): ovipositor length 63, gonostylus 10.5 
[mid tibia 52]. 

Male. Unknown. 

Hosts. Reared from the eggs of Cricula elaezia 
(Lepidoptera: Saturniidae). 

Distribution. Indonesia. 



36 



D. -W.HUANG AND J.S. NOYES 



Material examined. Type material. Holotype 
$, INDONESIA: Java, Pekalongen, Cinchona 
Plantation, ex Cricula elaezia, iii.1975 (M. Eddy) 
(BMNH). Paratypes: 1$, same data as holotype 
(BMNH). 

Comments. Ooencyrtus dione can be separated 
from other species of the genus by the broadly 
flattened scape, bicolored flagellum and enlarged 
clava. 

Ooencyrtus dry as sp.n. 

(Figs 106, 107) 

Diagnosis. Female: body blackish with a weak 
metallic sheen on head and dorsum of thorax; 
radicle blackish, rest of antennae whitish; all 
coxae and hind femora dark brown, blackish; 
remainder of legs white or yellow; forewing with 
a slightly oblique, median infuscate band (Fig. 
107); mandibles with one small tooth and broad 
truncation; antennae with F1-F5 distinctly longer 
than broad, F6 subquadrate or somewhat trans- 
verse; clava obliquely truncate at apex (similar to 
Fig. 16), with sensory area enlarged; frontover- 
tex one-quarter head width and narrowing ante- 
rior to ocelli; scutellum with regular, fine, 
punctate-reticulate sculpture, only its posterior 
margin smooth; postmarginal vein very short, 
nearly absent; ovipositor not or hardly exserted. 
Male: unknown. 

Female. Length about 0.85-1.05 mm (holotype 
1.02 mm). 

Frontovertex weakly metallic green; lower face 
including interantennal prominence and genae 
weakly metallic purple; antennae white except 
radicle blackish; sometimes clava yellowish; 
pronotum, mesoscutum, axillae and scutellum 
blackish with weak purplish tinge; scutellum dull, 
posterior margin smooth and purplish; 
mesopleuron and propodeum black-brown; 
coxae concolorous with mesopleuron, mid and 
fore femora white, hind femora totally blackish, 
all tibiae white, all tarsi yellow; forewing with an 
oblique, median infuscate band; gaster dark 
purple brown, basal tergites slightly metallic 
green or blue; ovipositor sheaths yellowish. 

Ocellar area with fine, regular, polygonal 
sculpture; sculpture on genae longitudinally 
elongate; ocelli forming an acute (about 50°) 
angle, posterior ocelli separated from occipital 
margin by about 2.5 times their own lengths and 
nearly touching eye margins; frontovertex one- 
quarter head width or less, the inner eye margins 
slightly convergent anterior to ocelli; eyes incon- 
spicuously hairy; occipital margin hardly 



rounded; head in facial view as broad as long; 
antennae inserted distinctly below the ventral 
eye margins, toruli separated from mouth margin 
by about their own lengths; clypeus with lower 
margin slightly protruding; antennae similar to 
Fig. 16, with pedicel plus flagellum about 1.3 
times head width; pedicel as long as following 
two segments together; Fl-5 distinctly longer 
than broad, F6 quadrate, or somewhat trans- 
verse; clava slightly longer than F4-6 combined, 
obliquely truncate at apex, with sensory area 
enlarged and extending along whole of truncate 
area, nearly half length of clava; mandibles 
robust, with one small tooth and a broad trunca- 
tion which is minutely denticulate. Relative mea- 
surements (holotype): head width 40, head 
height 40, minimum frontovertex width 10, POL 
4.5, OPL 7.5, OOL 0.5, eye length 27 eye width 
23, malar space 16, scape length 19, scape width 
3; other proportions of antennae similar to Fig. 
16. 

Mesoscutum with shallow, imbricate sculpture; 
scutellum with conspicuously deeper, fine 
punctate-reticulate sculpture (Fig. 106), only 
posterior margin smooth and shiny, and apex 
moderately pointed; forewing with venation and 
distribution of setae basally as in Fig. 107; costal 
cell narrow; linea calva closed posteriorly; post- 
marginal vein very short, almost absent. Relative 
measurements (holotype): forewing length 94, 
forewing width 38, hindwing length 65, hindwing 
width 14. 

Gaster about as long as thorax or shorter; 
ovipositor (similar to Fig. 335) more or less 
hidden; hypopygium similar to Fig. 36. Relative 
measurements (paratype): ovipositor length 83, 
gonostylus 17 [mid tibia 105]. 

Male. Unknown. 

Variation. Very little in material available, 
other than that described above under colour. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 450m, BMNH Canopy fog #5, 
2.ii.l985 (BMNH). Paratypes, INDONESIA: all 
same data as holotype but 1$, BMNH Canopy 
#3, 8.H.1985; 3$, BMNH Canopy #11, 
10.iii.1985; 3$ iv.1985 (J.S. Noyes); 1$, v.1985 
(J. Martin); 2$, BMNH Canopy #13, 
ll.vii.1985; 1$, BMNH Canopy fog #15, 
19.vii.1985. Material in BMNH, IZAS, MZB. 

Comments. Females of Ooencyrtus dryas can de 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



37 



distinguished from other species of the genus by 
the almost totally white antennae and legs con- 
trasting with the rest of the body, infuscate 
forewings, obliquely truncate clava and eyes con- 
verging anterior to the ocelli. 

Ooencyrtus ceres sp.n. 

(Figs 108-112) 

Diagnosis. Body blackish, head and dorsum of 
thorax dorsally metallic blue or green; scutellum 
more strongly metallic than mesoscutum; all 
coxae black, femora usually brown on sides, 
otherwise yellow; tibiae yellow; mandibles with 
one tooth and a broad truncation; anterior three- 
fifths of scutellum with regular, fine punctate- 
reticulate sculpture, posterior part smooth and 
shiny (Fig. 109). Female: antennae (Fig. 108) 
with funicle dark testaceous and clava yellow; 
frontovertex less than one-quarter head width; 
all funicle segments longer than broad; clava 
hardly broader than funicle segments, its sutures 
parallel and sensory area small and at apex only; 
forewing (Fig. 110) with linea calva closed poste- 
riorly; ovipositor not exserted or hardly so. 
Male: similar to female but antennae unicolorous 
testaceous-yellow; all funicle segments at least 
twice as long as broad and clothed in setae about 
twice as long as diameter of segments; frontover- 
tex about one-third head width. 

Female. Length about 0.8-1.05 mm (holotype 
0.8 mm). 

Head blackish with a green or purple sheen on 
frontovertex; the face, interantennal prominence 
and genae green, more metallic; lowest margins 
of toruli connected by a very narrow, broadly 
u-shaped line; antennae with radicle, dorsal sides 
of pedicel and funicle segments dark testaceous, 
ventral sides of pedicel and funicle segments 
testaceous, scape and clava yellow; pronotum, 
mesoscutum and axillae blackish with a metallic 
green or blue sheen; scutellum metallic green 
mixed with some blue, clearly brighter than 
mesoscutum, posterior two-fifths very shiny and 
with a purplish sheen; mesopleuron and propo- 
deum dark brown; coxae, dorsal and ventral 
sides of femora blackish, trochanters, apices of 
femora, tibiae and tarsi yellow, pretarsi brown; 
wings hyaline, venation testaceous; gaster black- 
ish with a slight metallic green sheen dorsally; 
ovipositor sheaths dark brown. 

Head in ocellar area with fine, shallow, 
polygonally reticulate sculpture; reticulate sculp- 
ture on genae longitudinally elongate; ocelli 
forming an angle of about 60°, posterior ocelli 
separated from occipital margin by about 2 times 



their own length and almost touching eye mar- 
gins; frontovertex less than one-quarter head 
width; eyes not conspicuously hairy; occipital 
margin hardly rounded; antennae inserted 
slightly below the ventral eye margins (Fig. Ill), 
toruli separated from mouth margin by more 
than their own lengths; clypeal margin slightly 
produced medially (Fig. Ill); antennae (Fig. 
108) with all funicle segments conspicuously 
longer than broad, clava about as long as F4-6 
together, sutures parallel and sensory area at 
extreme apex only; mandibles (similar to Fig. 
333) with one teeth and a broad, minutely den- 
ticulate truncation. Relative measurements 
(holotype): head width 41, head height 36, mini- 
mum frontovertex width 10, POL 4, OPL 5, 
OOL 0.5, eye length 26, eye width 21, malar 
space 15, scape length 19, scape width 4.5; other 
proportions of antenna as in Fig. 108. 

Mesoscutum with shallow, imbricate sculpture; 
scutellum (Fig. 109) with conspicuously deeper, 
fine, punctate-reticulate sculpture in anterior 
three-fifths, posterior two-fifths smooth and very 
shiny; fore wing with venation and distribution of 
setae basally as in Fig. 110; linea calva closed 
posteriorly. Relative measurements (holotype): 
forewing length 100, forewing width 38; hind- 
wing length 65; hindwing width 15. 

Gaster shorter than thorax; ovipositor (similar 
to Fig. 335) not or hardly exserted, hypopygium 
as in Fig. 112. Relative measurements 
(paratype): ovipositor length 102, gonostylus 19 
[mid tibia 95]. 

Male. Length: 0.85 mm. Generally very similar 
to female but for antennae, wider frontovertex, 
coloration of legs and structure of genitalia. 
Antennae testaceous-yellow, slightly darker 
proximally; mid-femora entirely yellow; fron- 
tovertex about one-third head width; antennae 
with pedicel plus flagellum about 1.5 times as 
long as head width; Fl, F5 and F6 equal in 
length, subequal to pedicel, each 2 times as long 
as broad; F2-4 subequal in length and slightly 
longer than other segments; clava about 3.5 
times as long as broad, as long as two proceeding 
segments together; funicle clothed in setae about 
2 times as long as diameter of segments. 

Variation. Very little in material available. 

Hosts. Reared from eggs of Cratoplatys sp. 
(Heteroptera: Plataspidae). 

Distribution. Brunei, Malaysia. 

Material examined. Type material. Holotype 
$, MALAYSIA: Sabah, Tawau, ex eggs of 
Cratoplatys sp., CIE A18090, ll.vi.1986 



38 



D. -W.HUANG AND J.S. NOYES 



(G.T.Lim) (BMNH). Paratypes, MALAYSIA: 
1$, lCf, same data as holotype; BRUNEI, 2$, 
Bukit Sulang, Nr. Lamunin, B.M. 1982-388, 
20.viii-10.ix.1982 (N.E. Stork) (BMNH). 

Comments. Females of Ooencyrtus ceres can be 
separated from other species of the genus by the 
unique combination of a bicolorous flagellum, 
finely punctate-reticulate scutellum which is 
shiny in posterior two-fifths, filiform antennae 
with all funicle segments longer than broad and 
generally yellow legs with femora normally mar- 
gined brown (see also comments under ilion, iris, 
elissa and bacchus below). 

Ooencyrtus ilion sp.n. 

(Figs 113-116) 

Diagnosis. Female: body generally blackish, 
but head slightly tinged with purple and green, 
mesoscutum metallic blue, scutellum brighter 
green; antennae generally testaceous; all coxae 
blackish, hind femora and fore femora mainly 
dark brown except at apices; mid femora yellow; 
mandible (Fig. 114) with one very small tooth 
and a broad, minutely denticulate truncation; 
flagellum clavate (Fig. 113), all funicle segments 
subquadrate, Fl-3 conspicuously smaller than 
F4-6; clava much broader than F6, sutures 
oblique and sensory area extending along ventral 
part of apical segment, about half as long as 
clava; frontovertex one-sixth head width; scutel- 
lum anteriorly finely punctate-reticulate, poste- 
rior one-third or less smooth and shiny; forewing 
(Fig. 115) with conspicuous basal naked area, 
linea calva open posteriorly; postmarginal vein 
about two-third of stigmal vein; ovipositor not 
exserted. 

Female. Length 0.9-1.0 mm (holotype 0.95 
mm). 

Head blackish, weakly metallic green, ocellar 
area slightly purple; face metallic green, interan- 
tennal prominence dorsally purple; antennae 
with radicle dark brown, scape yellow but 
marked with brown basally; pedicel mainly 
brownish but apex testaceous, flagellum 
testaceous-yellow, Fl-3 and clava slightly 
darker; pronotum black, weakly metallic; mesos- 
cutum and axillae blackish with strong blue or 
blue green metallic sheen; scutellum green, with 
blue reflections, apically blue, very shiny; 
mesopleuron and propodeum dark brown; coxae 
concolorous with mesopleuron, fore-femora and 
hind-femora mainly dark brown except for yel- 
low apices; mid-femora totally yellow, rest of 
legs yellow; wings hyaline, venation testaceous; 



gaster black, slightly metallic green basally; ovi- 
positor sheaths dark brown. 

Head with very shallow, fine, polygonally 
reticulate sculpture in ocellar area; sculpture on 
genae longitudinally elongate; ocelli forming an 
acute angle of about 40°, posterior ocelli sepa- 
rated from occipital margin by about 1.5 times 
their own lengths and touching eyes; frontover- 
tex about one-sixth head width; eyes not con- 
spicuously hairy; occipital margin hardly 
rounded; antennae inserted distinctly below ven- 
tral eye margins, toruli separated from mouth 
margin by less than their own lengths; antennae 
(Fig. 113) with pedicel plus flagellum about 0.9 
times head width; all funicle segments subquad- 
rate, Fl-3 somewhat transverse and distinctly 
smaller than following segments F4-6; clava con- 
spicuously broader than F6, about as long as 
F3-F6 together, with outer suture moderately 
oblique, sensory area and about half as long as 
clava; mandibles (Fig. 114) robust, with one 
small tooth and a broad inner, minutely denticu- 
late truncation. Relative measurements (holo- 
type): head width 47, head height 42, minimum 
frontovertex width 8, POL 3, OPL 4, OOL 0, eye 
length 30 eye width 28, malar space 17, scape 
length 18, scape width 4.5; other proportions of 
antennae as in Fig. 113. 

Mesoscutum with very shallow polygonally 
reticulate or imbricate sculpture; scutellum with 
distinctly deeper, punctate-reticulate sculpture, 
shallower in apical one-third and with posterior 
one-fifth smooth, apex moderately pointed; 
forewing with venation and distribution of setae 
basally as in Fig. 115, linea calva open posteri- 
orly. Relative measurements (holotype): forew- 
ing length 99, forewing width 41. 

Gaster about two-thirds as long as thorax; 
ovipositor (Fig. 116) not exserted, with gonosty- 
lus only about one-sixth as long as ovipositor; 
hypopygium similar to Fig. 36. Relative measure- 
ments (paratype): ovipositor length 123, gono- 
stylus20[midtibia96]. 

Male. Unknown. 

Variation. Very little in material available. 

Hosts. Reared from the eggs of Amblypelta 
cocophaga (Hemiptera: Coreidae) from coconut. 

Distribution. Solomon Islands. 

Material examined. Type material. Holotype 
?, SOLOMON ISLANDS: Lunga, Guadalca- 
nal, ex Amblypelta cocophaga eggs from coco- 
nut, ll.x.1937 (J.D.D. Love) (BMNH). 
Paratypes, SOLOMON ISLANDS: 5$, same 
data as holotype; 3$, Baunami, ex eggs of 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



39 



Amblypelta cocophaga, 23. vi. 1937 (J.S. Phillips). 
Material in BMNH, IZAS. 

Comments. Ooencyrtus ilion may be closest to 
ceres, both species having very similar habitus, 
general coloration and sculpture on the scutel- 
lum. O. ilion can be distinguished from ceres by 
having a clavate flagellum with very short seg- 
ments and clava with an enlarged, oblique sen- 
sory area whilst in ilion the flagellum is filiform 
with elongate segments and the clava has a small 
sensory area at extreme apex only. 

O. ilion is also superficially similar to iulus (see 
comments under iulus below). 

Ooencyrtus iris sp.n. 

(Figs 117-119) 

Diagnosis. Female: body generally dark brown 
or blackish, face metallic green, frontovertex 
slightly coppery; mesoscutum dull but slightly 
metallic green, scutellum anteriorly dull pur- 
plish, posterior part and sides metallic blue or 
green; antennae testaceous; all coxae dark 
brown, legs yellow but with fore and hind femora 
usually margined dark brown; mandibles with a 
small tooth and a broad, slightly convex, 
minutely denticulate, truncation; flagellum fili- 
form, all funicle segments longer than broad; 
clava hardly broader than funicle, sutures paral- 
lel, sensory area at extreme apex only; scutellum 
anteriorly with shallow sculpture, posterior half 
more or less smooth and shiny; forewing (Fig. 
119) without conspicuous basal naked area; linea 
calva closed posteriorly; ovipositor not exserted. 
Male: unknown. 

Female. Length about 0.85-1.05 mm (holotype 
1.0 mm). 

Head in ocellar area dull with a slight green 
and coppery sheen; face, brighter metallic green, 
sometimes bluish or brassy; a very thin metallic 
purple v-shaped line connects lowest margins of 
antennal toruli; antennae testaceous-yellow; 
pronotum, mesoscutum, axillae and anterior half 
of scutellum blackish with weak metallic purple 
or green sheen; posterior half of scutellum con- 
spicuously metallic blue, green and purple, much 
brighter than anterior half; mesopleuron and 
propodeum dark brown; coxae concolorous with 
mesopleuron, legs yellow but with fore and hind 
femora margined dorsally and ventrally brown; 
wings hyaline, venation testaceous; gaster black- 
ish with a slight metallic green sheen; ovipositor 
sheaths yellow. 

Head in ocellar area with shallow, regular, 
polygonally reticulate sculpture, that on genae 



longitudinally elongate; ocelli forming an acute 
angle of about 50°, posterior ocelli separated 
from occipital margin by about 1.5-2 times their 
own lengths and almost touching eye margins; 
frontovertex one-fifth head width; eyes not con- 
spicuously hairy; occipital margin slightly 
rounded; antennae inserted just below the ven- 
tral eye margin (Fig. 117); toruli separated from 
mouth margin by little more than their own 
lengths; clypeal margin slightly protuberant 
medially (Fig. 117); antennae (similar to Fig. 
108) with pedicel plus flagellum 1.1 times as long 
as head width; all funicle segments conspicuously 
longer than broad, clava about as long as F4-6 
together, with sutures parallel and sensory area 
at extreme apex only; mandibles (similar to Fig. 
333) with one tooth and a slightly convex, broad, 
minutely denticulate truncation. Relative mea- 
surements (holotype): head width 41, head 
height 38, minimum frontovertex width 9, POL 
4, OPL 4, OOL 0.3, eye length 28, eye width 23, 
malar space 15, scape length 17, scape width 3.5; 
other proportions of antenna similar to Fig. 108. 

Mesoscutum with shallow, imbricate sculpture; 
scutellum basally with similar, but finer sculp- 
ture, posterior half and sides smooth, its apex 
rounded; fore wing with venation and distribu- 
tion of setae basally as in Fig. 119. Relative 
measurements (holotype): forewing length 93, 
forewing width 37; hindwing length 60; hindwing 
width 14. 

Gaster about as long as thorax, ovipositor 
(similar to Fig. 335) not exserted, with gonosty- 
lus about one-sixth as long as ovipositor; hypopy- 
gium as in Fig. 118. Relative measurements 
(paratype): ovipositor length 90, gonostylus 15 
[mid tibia 90]. 

Male. Unknown. 

Variation. Very little in material available. 

Hosts. Unknown. 

Distribution. Papua New Guinea. 

Material examined. Type material. Holotype 
$, PAPUA NEW GUINEA: Madang Prov., 
Laing, 20.vi.1982 (P. Grootaert) (BMNH). 
Paratypes, PAPUA NEW GUINEA: 1$, same 
data as holotype; 1$, same data but vii.1982; 
39, same data but xi.1982. Material in BMNH. 

COMMENTS. Ooencyrtus iris is probably close to 
Ooencyrtus ceres, both species having a very 
narrow purple line connecting the lowest margins 
of the toruli, more or less filiform flagellum, 
scutellum smooth posteriorly and linea calva 
closed posteriorly. O. iris differs from ceres in 



40 



D. -W.HUANG AND J.S. NOYES 



having a unicolorous flagellum, anterior part of 
scutellum with relatively shallow imbricate sculp- 
ture and ovipositor sheaths yellow. In ceres the 
flagellum is bicolorous, the anterior part of the 
scutellum has relatively deep punctate-reticulate 
sculpture, and the ovipositor sheaths are dark 
brown. 

The species is also close to Ooencyrtus elissa 
and bacchus (see comments under elissa and 
bacchus below). 

Ooencyrtus javanicus Mercet 

(Figs 120-124) 

Ooencyrtus (Schedius) javanicus Mercet, 1922a: 
152-153. Lectotype $ (designated by Noyes, 
1981), Indonesia (IEEM, examined). 

Ooencyrtus javanicus Mercet; Ferriere, 1931: 

282. 

Diagnosis. Female (length about 0.8-1 mm): 
body generally dark brown, metallic green or 
blue on head, dorsum of thorax and gaster; 
apical half and sides of scutellum more strongly 
metallic blue or purplish; antennae testaceous- 
yellow; coxae and fore-femora mostly blackish, 
rest of legs yellow; tegulae dark brown; oviposi- 
tor sheaths yellow; mandibles (Fig. 121) with a 
small tooth and a broad truncation; frontovertex 
about one-quarter head width; antennae (Fig. 
120) with funicle segments at most a little longer 
than broad, F6 distinctly transverse; clava with 
sensory area at apex only; anterior half of scutel- 
lum with similar shallow sculpture to mesoscu- 
tum, this becoming shallower towards apex so 
that posterior half is smooth; forewing (Fig. 123) 
with conspicuous naked area basally; ovipositor 
(Fig. 122) only slightly exserted, slightly longer 
than mid tibia; hypopygium similar to Fig. 36. 
Male (length 0.6-1.0 mm): similar to female but 
antennae and legs, including coxae, more or less 
completely yellow; bases of tegulae yellow; 
antennae (Fig. 124) with all funicle segments at 
least about twice as long as broad and clothed in 
setae about twice as long as diameter of seg- 
ments; aedeagus about two-thirds as long as mid 
tibia. 

Hosts. Recorded from the eggs of Lepidoptera 
by Mercet (1922a) and recorded below from the 
eggs of Cricula sp. (Lepidoptera: Saturniidae). 

Distribution. Indonesia. 

Material examined. Type material. Lecto- 
type $ : no data other than 'Schedius javanicus'' 
[according to Mercet the specimen came from 
Indonesia, Java and was reared as an endopha- 



gous parasite of the eggs of a lepidopteran] 
(IEEM). 

Other material. INDONESIA: 4$, 5cf, ex 
eggs of Cricula, CIE A2125, vi.1990. In BMNH. 

Comments. In general habitus and coloration, 
especially of the legs, Ooencyrtus javanicus is 
extremely similar to O. iulus. The two species 
can be separated on the relative proportions of 
the marginal, postmarginal and stigmal veins and 
relatively more densely setose basal part of 
forewing in iulus (compare Figs 123 and 128), the 
yellow ovipositor sheaths of javanicus (brown in 
iulus) and ovipositor longer than mid tibia in 
javanicus (not longer than mid tibia in iulus) with 
longer, more slender gonostyli. 

Ooencyrtus iulus sp.n. 

(Figs 125-128) 

Diagnosis. Female: body blackish weakly pur- 
plish or green on head and pronotum, mesoscu- 
tum quite strongly metallic green, blue or purple, 
scutellum largely blue or purple; antennae with 
radicle black, remainder yellow; all coxae black, 
fore-femora black, rest of legs mostly yellow; 
mandibles with one small tooth and a broad, 
minutely denticulate truncation; frontovertex not 
more than about one-quarter head width (Fig. 
127); flagellum slightly clavate; Fl^ subquad- 
rate; clava hardly broader than F6, with sensory 
area slightly enlarged, but more or less restricted 
to apex only; scutellum anteriorly with shallow 
sculpture as mesoscutum, posteriorly smooth and 
shiny; forewing (Figs 128) without conspicuous 
basal naked area; linea calva open posteriorly; 
ovipositor not exserted. Male: coloration similar 
to female, but dorsum of thorax much duller, 
coppery purple; antennae with all funicle seg- 
ments at least twice as long as broad and clothed 
in setae more than 2 times as long as diameter of 
segments; frontovertex slightly wider than one- 
third head width. 

Female. Length about 0.7-0.9 mm (holotype 
0.90 mm). 

Head blackish with weak metallic green or 
purplish sheen on frontovertex; face purplish; at 
least top of scrobes metallic green, but some- 
times also rest of face and genae; antennae with 
radicle black, other segments yellow; pronotum, 
purplish brown; mesoscutum quite strong metal- 
lic green or blue margined purple, anterior part 
of scutellum relatively dull purplish, posterior 
part strong metallic green or blue; mesopleuron 
dark purple-brown; coxae concolorous with 
mesopleuron, fore-femora almost totally dark 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



41 



brown, rest of legs yellow but hind femora some- 
times with a black mark along dorsal margin; 
wings hyaline, venation testaceous; gaster black 
with a metallic green or blue sheen; ovipositor 
sheaths black. 

Head with fine, shallow, polygonally reticulate 
sculpture on frontovertex; sculpture on genae 
longitudinally elongate; ocelli forming an equi- 
lateral triangle, the posterior ocelli separated 
from occipital margin by about one and half 
times their own lengths and almost touching eye 
margins; frontovertex one-fifth to nearly one- 
quarter head width (Fig. 127); eyes not conspicu- 
ously hairy; occipital margin hardly rounded; 
antennae inserted slightly below ventral eye mar- 
gin, toruli separated from mouth margin by less 
than their own lengths; antennae (Figs 125, 126) 
with pedicel plus flagellum about 0.8 times head 
width; ¥\-A subquadrate, F6, sometimes includ- 
ing F5 somewhat transverse; clava not conspicu- 
ously broader than F6, slightly longer than F4-6 
combined with sensory area slightly enlarged and 
about half as long as apical segment; mandibles 
(similar to Fig. 139) with a small tooth and a 
broad, minutely denticulate truncation. Relative 
measurements (holotype): head width 48, head 
height 40, minimum frontovertex width 9, POL 
5, OPL 4, OOL 0.3, eye length 28 eye width 24, 
malar space 15, scape length 18, scape width 4; 
other proportions of antennae as in Figs 125 and 
126. 

Mesoscutum with shallow imbricate sculpture; 
anterior part of scutellum with similar sculpture, 
posterior half smooth and shiny; forewing with 
venation and distribution of setae basally as in 
Fig. 128, without conspicuous naked area 
basally; linea calva open posteriorly. Relative 
measurements (holotype): forewing length 90, 
forewing width 38. 

Gaster conspicuously shorter than thorax, in 
dorsal view nearly rounded apically; ovipositor 
hidden; ovipositor similar to Fig. 335; hypopy- 
gium similar to Fig. 36. Relative measurements 
(paratype): ovipositor length 72, gonostylus 13 
[mid tibia 73]. 

Male. Length 0.7 mm. 

Generally similar to female but for coloration 
of dorsum of thorax, frontovertex width and 
structure of antennae and genitalia. Thorax dor- 
sally generally slightly metallic with a coppery or 
brassy sheen; apex of scutellum metallic blue- 
green; frontovertex a little more than one-third 
head width with posterior ocelli distinctly sepa- 
rated from eye margin; antennae with pedicel 
plus flagellum about 1.5 times head width; all 
funicle segments at least 2 times as long as broad, 



with setae at least 2 times as long as diameter of 
segments, clava as F5-6 together. 

Variation. In the female the metallic sheen on 
the head and thorax varies in both intensity and 
coloration as described above; the frontovertex 
varies from about one-fifth to one-quarter head 
width; antennae may have ¥\-A distinctly longer 
than broad (Fig. 126), or Fl-2 subquadrate (Fig. 
125). 

Hosts. Recorded from the eggs of Eocanth- 
econa furcellata (Hemiptera: Pentatomidae). 

Distribution. India. 

Material examined. Type material. Holotype 
$, INDIA: Madras, ex egg Eocanthecona furcel- 
lata, CIE A 19990 (1988) (BMNH). Paratypes, 
INDIA: 4$, ld\ same data as holotype; 29, 
Tamil Nadu, 3 km. E. Manjaler Dam, 
15-18.X.1979 (J.S. Noyes); 19, Tamil Nadu, 
Mangarai Forest, 28. ix. 1979 (J.S. Noyes); 19, 
Karnataka, Bannerghatta N.P., 5.xi.l979 (Z. 
Boucek & J.S. Noyes). Material in BMNH. 

Comments. Females of Ooencyrtus iulus are 
similar to those of ilion in general habitus and 
coloration. The two species can be separated on 
the sculpture of the scutellum, the scutellum of 
ilion being punctate-reticulate basally and much 
deeper than that on mesoscutum, whilst that of 
iulus is relatively shallow and not deeper than 
that on mesoscutum. There is also a marked 
difference in the structure of the clava with iulus 
having only a relatively small sensory area on the 
clava which is restricted to the apex only, whilst 
that of ilion is very nearly half as long as the clava 
itself. 

O. iulus is also very similar to javanicus (see 
comments under javanicus above). 

Ooencyrtus ixion sp. n. 

(Figs 129-132) 

Diagnosis. Female: body generally blackish; 
head weakly metallic blue-green, dorsum of tho- 
rax dull metallic blue-green; antennae mostly 
testaceous; legs with all coxae dark brown and 
legs mostly yellow but with hind femora and 
sometimes fore femora proximally testaceous- 
brown; frontovertex about one-third head width; 
mandibles with two teeth and a truncation, sec- 
ond tooth very short, almost absent; scutellum 
with sculpture similar to that on mesoscutum, 
towards apex almost smooth; apex of scutellum 
pointed; flagellum weakly clavate, almost fili- 
form with all funicle segments only slightly 
longer than broad; clava slightly broader than 



42 



D. -W.HUANG AND J.S. NOYES 



F6, its sutures parallel and sensory area small and 
at apex only; forewing (Fig. 130) with conspicu- 
ous naked basal area; linea calva open posteri- 
orly; postmarginal vein very short, almost 
absent; ovipositor not or hardly exserted. Male: 
similar to female, but frontovertex a little more 
than one-third head width; antennae with all 
funicle segments subequal and slightly longer 
than broad and clothed with setae that are about 
as long as diameter of segments or less. 

Female. Length about 8.0-9.5 mm (holotype 
0.9 mm). 

Head blackish, weakly metallic green on fron- 
tovertex, face metallic green, dorsal part of 
scrobes and interantennal prominence purple; 
antennae with radicle brownish, scape with 
proximal half yellow and distal half brownish, 
pedicel mainly brownish but tip testaceous, fla- 
gellum testaceous; pronotum, mesoscutum, axil- 
lae and scutellum blackish with weak metallic 
blue-green lustre; mesopleuron and propodeum 
blackish; coxae concolorous with mesopleuron, 
fore femora yellowish, sometimes proximally 
brownish, hind femora with proximal two-thirds 
brown and distal one-third or less yellow, rest of 
legs yellow; wings hyaline, venation testaceous; 
gaster black with metallic purple tinge; oviposi- 
tor sheaths black. 

Head with regular, fine, polygonally reticulate 
sculpture on frontovertex; sculpture on genae 
longitudinally elongate; ocelli relatively large 
and forming a right angle, posterior ocelli sepa- 
rated from occipital margin and eye margins by a 
little less than their own lengths; frontovertex 
about one-third head width; eyes not conspicu- 
ously hairy; occipital margin not distinctly 
rounded; head with genae strongly convergent 
towards mouth (Fig. 132); antennae inserted 
distinctly below ventral eye margin, toruli sepa- 
rated from mouth margin of face by about their 
own lengths; antennae (Fig. 129) with pedicel 
plus flagellum about as long as head width; all 
funicle segments only slightly longer than broad, 
clava slightly broader than F6, about as long as 
F4-6 together, sensory area at apex only; man- 
dibles (similar to Fig. 143) with one tooth and a 
broad truncation. Relative measurements (holo- 
type): head width 42, head height 37, minimum 
frontovertex width 14, POL 9, OPL 2, OOL 1.5, 
eye length 25 eye width 20, malar space 15, scape 
length 18, scape width 3.5; other proportions of 
antennae as in Fig. 129. 

Mesoscutum with shallow imbricate sculpture; 
scutellum with similar sculpture basally, but 
becoming gradually shallower apically so that 
apical one-third or so is smooth; apex pointed; 



forewing with venation and distribution of setae 
as in Fig. 130, with a distinct naked basal area; 
linea calva open posteriorly; postmarginal vein 
very short, almost absent. Relative measure- 
ments (holotype): forewing length 92, forewing 
width 40; hindwing length 57; hindwing width 14. 
Gaster conspicuously shorter than thorax; ovi- 
positor (Fig. 131) not exserted; hypopygium 
similar to Fig. 25. Relative measurements 
(paratype): ovipositor length 123, gonostylus 20 
[mid tibia 96]. 

Male. Length 0.79 mm. Generally similar to 
female but differs in relative width of frontover- 
tex, antennal structure and genitalia. Frontover- 
tex a little more than one-third head width; ocelli 
forming an obtuse angle of about 100°; posterior 
ocelli separated from occipital margin by about 
half of their own length; antennae with pedicel 
plus flagellum about 1.25 times head width; 
pedicel and all funicle segments subequal and 
slightly longer than broad, clava as long as F5-6; 
funicle segments clothed in setae that are about 
as long as diameter of segments or less. 

Variation. Female with fore-femora some- 
times totally yellow. 

Hosts. Reared from the eggs of a myrmeleontid 
on grass (Neuroptera: Myrmeleontidae). 

Distribution. India. 

Material examined. Type material. Holotype 
$, INDIA: Tamil Nadu, Padappai, ex eggs of 
Myrmeleontidae from grass, CIE A20314, 
18.iii.1989 (BMNH). Paratypes, INDIA: 50$, 
lcf, same data as holotype (only 4 females 
mounted, remainder in poor condition and 
stored in a gelatin capsule). Material in BMNH. 

Comments. Females of Ooencyrtus ixion can be 
separated from other species of the genus by the 
frontovertex being about one-third head width, 
the posterior ocelli well separated from the eye 
margin and the virtually absent postmarginal 
vein of the forewing. It is superficially similar to 
O. ferrierei and the extralimital O. pityocampae. 
However both ferrierei and pityocampae have the 
coxae and legs totally yellow, and in ferrierei the 
gaster is largely yellow whilst in pityocampae the 
funicle segments are all clearly longer than broad 
and relatively much longer than ixion. 

Ooencyrtus elissa sp.n. 

(Figs 133-137) 

Diagnosis. Female. Body generally dark brown 
or blackish with head metallic green, meoscutum 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



43 



weakly metallic blue-green and apical two-thirds 
of scutellum metallic green; antennae bicolorous, 
mostly testaceous-brown but clava yellow; fore 
and mid legs, including coxae, yellow; hind- 
coxae black, hind legs mainly yellow; ovipositor 
with gonostylus pale yellow; mandibles (Fig. 137) 
with one small tooth and a broad, minutely 
denticulate truncation; flagellum filiform, Fl-2 
subquadrate, F3-6 larger and longer than broad; 
clava slightly broader than F6, with sutures sub- 
parallel and sensory part at apex only; anterior 
half of scutellum with sculpture very slightly 
deeper than that on meoscutum, remainder 
smooth; forewing (Fig. 135) with only a small 
naked basal area; frontovertex a little less than 
one-third head width; ovipositor not exserted. 

Male. Unknown. 

Female. Length about 1.08 mm (holotype). 

Head blackish with a strong metallic green 
sheen, weaker in ocellar area, slightly brassy or 
bluish in scrobal area and a thin purple, u-shaped 
line connecting lowest margins of toruli; anten- 
nae with radicle black-brown, scape mainly yel- 
low but narrowly margined brown; pedicel 
proximally brown, distally testaceous; funicle 
segments testaceous-brown, F4-6 ventrally yel- 
lowish; clava yellow, first segment brownish 
basally; pronotum, axillae and anterior one-third 
of scutellum black with a purple tinge; mesoscu- 
tum metallic blue; posterior half of scutellum 
shining metallic green; mesopleuron and propo- 
deum dark brown; fore and mid coxae yellow, 
hind coxae dark brown; legs yellow but for a 
narrow black line along dorsal part of hind 
femora; wings hyaline, venation testaceous; 
gaster dark purple-brown with weak blue tinge 
basally; ovipositor with sheaths pale yellow. 

Frontovertex with regular, raised polygonal 
sculpture, deeper in ocellar area; sculpture on 
lower face and genae longitudinally elongate; 
ocelli forming a right angle; posterior ocelli sepa- 
rated from occipital margin by about 1.5 times 
their own lengths and almost touching eye mar- 
gins; frontovertex a little less than one-third head 
width; eyes not conspicuously hairy; occipital 
margin rounded; antennae inserted distinctly 
below ventral eye margins; toruli separated from 
mouth margin by slightly more than their own 
lengths; clypeus slightly produced medially; 
antennae (Fig. 134) with pedicel plus flagellum as 
long as head width; Fl-2 subquadrate, F3-6 
longer than broad, subequal in length; clava only 
slightly broader than F6, about as long as F4-6 
together, with sutures parallel and sensory part 
small and at apex only; mandibles (Fig. 137) with 



one small tooth and a broad, minutely denticu- 
late truncation. Relative measurements: (holo- 
type): head width 41, head height 36, minimum 
frontovertex width 12, POL 6, OPL 5, OOL 0.5, 
malar space 16, eye length 25, eye width 20, 
scape length 18, scape width 3.5; other propor- 
tions of antenna as in Fig. 134. 

Mesoscutum with shallow imbricate sculpture; 
scutellum anteriorly with hardly deeper, more 
polygonal sculpture becoming gradually shal- 
lower so that posterior half is almost totally 
smooth and shiny; apex hardly pointed; fore 
wing with venation and distribution of setae 
basally as in Fig. 135, with only a small naked 
basal area; linea calva conspicuously open poste- 
riorly. Relative measurements (holotype): forew- 
ing length 95, forewing width 40; hindwing length 
64; hindwing width 14. 

Gaster about as long as thorax; ovipositor 
hidden, similar to Fig. 335 with gonostylus about 
one-sixth as long as ovipositor; hypopygium as in 
Fig. 136. Relative measurements (paratype): ovi- 
positor length 81, gonostylus 14 [mid tibia 80]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Thailand. 

Material examined. Type material. Holotype 
9, THAILAND: Chieng Mai, vii.1984 (D. Jack- 
son) (BMNH). Paratype, THAILAND: 1$, 
22.vii-13.viii.1984, other data same as holotype. 
Material in BMNH. 

Comments. Ooencyrtus elissa is closest to Ooen- 
cyrtus ceres, iris, and bacchus, all four species 
having the ventral margins of the toruli con- 
nected by a faint, thin purplish v-shaped or 
u-shaped line, similar mandibular structure, the 
clypeal margin slightly produced medially and 
the legs largely yellow sometimes with the hind 
femur dorsally margined blackish. Apart from 
the characters which separate these species in the 
key, elissa can be distinguished from ceres by the 
relatively shallower sculpture at the base of the 
scutellum (in ceres it is relatively deep and 
punctate-reticulate), relatively shorter funicle 
segments (in ceres all at least about 1.5 times as 
long as broad) and yellow ovipositor sheaths 
(blackish in ceres). O. elissa differs from iris in 
having a bicolorous flagellum (unicolorous testa- 
ceous in iris) and relatively shorter funicle seg- 
ments (all clearly longer than broad in iris). See 
also comments under bacchus and pilosus 
(below). 

Females of elissa are superficially similar to 
egeria but have the ovipositor sheaths pale yel- 



44 



D. -W.HUANG AND J.S. NOYES 



low, antenna with Fl-2 subquadrate, forewing 
with a relatively elongate postmarginal vein and 
regular, polygonally reticulate sculpture on the 
anterior part of the scutellum. O. egeria has the 
ovipositor sheaths black, antenna with Fl-2 dis- 
tinctly longer than broad, the postmarginal vein 
virtually absent and longitudinally elongate 
sculpture on the scutellum anteriorly. O. elissa is 
also similar to the extralimital O. dipterae (Ris- 
bec) but can be separated by the coloration of 
legs, density of setae at the base of the forewing 
and relative proportions of the antennal seg- 
ments. 

Ooencyrtus egeria sp.n. 

(Figs 138-140) 

Diagnosis. Body generally blackish, head and 
dorsum of thorax with a weak purple sheen, apex 
of scutellum metallic green; antennae except 
radicle yellow-testaceous; fore coxae yellow, 
other coxae dark brown; hind femora mainly 
brown, rest of legs yellow; ovipositor sheaths 
black; mandibles with one small tooth and a 
broad truncation; flagellum filiform; all funicle 
segments longer than broad; clava not or hardly 
broader than F6, the sutures parallel with sen- 
sory area slightly enlarged but at apex only; 
forewing (Fig. 140) with setae very fine, costal 
cell almost naked; postmarginal vein very short, 
almost absent; scutellum with longitudinally 
elongate reticulate sculpture anteriorly, and 
smooth in posterior one-quarter; apex moder- 
ately rounded; frontovertex a little less than 
one-third head width; ovipositor not exserted. 

Female. Length about 1.10 mm (holotype). 

Head blackish with weak purple sheen; anten- 
nae with radicle dark brown, the remainder 
testaceous-yellow; pronotum, mesoscutum, axil- 
lae and anterior three-fourths of scutellum black- 
ish with a weak purple tinge; posterior one- 
quarter of scutellum metallic green and much 
brighter than anterior part; mesopleuron and 
propodeum blackish with a purple sheen; legs 
with mid and hind coxae dark brown, hind 
femora brown and remainder yellow; wings hya- 
line, venation testaceous; gaster black with a 
weak blue or purple sheen; ovipositor sheaths 
black. 

Head with fine, regular, polygonally reticulate 
sculpture on frontovertex; sculpture on lower 
face and genae longitudinally elongate; ocelli 
forming an equilateral triangle; posterior ocelli 
separated from occipital margin by about their 
own lengths and almost touching eye margins; 
frontovertex a little less than one-third head 



width; eyes not conspicuously hairy; occipital 
margin rounded; antennae inserted below the 
ventral eye margins; toruli separated from mouth 
margin by about their own lengths; antennae 
(similar to Fig. 148) with pedicel plus flagellum 
as long as head width; all funicle segments con- 
spicuously longer than broad, subequal in length; 
clava as broad as funicle segments, about as long 
as F4-6, with sutures parallel and sensory part 
slightly enlarged but only at extreme apex; man- 
dibles (Fig. 139) with one small tooth and a 
broad straight truncation. Relative measure- 
ments (holotype): head width 40, head height 35, 
minimum frontovertex width 12, POL 6, OPL 
3.5, OOL 0.5, eye length 25, eye width 20, scape 
length 17, scape width 2.5; other proportions of 
antennae similar to Fig. 148. 

Mesoscutum with shallow, imbricate, almost 
polygonal sculpture; scutellum with very slightly 
deeper, longitudinally elongate sculpture in ante- 
rior three-quarters, this gradually becoming shal- 
lower posteriorly so the posterior quarter smooth 
and shiny; apex rounded; fore wing with vena- 
tion and distribution of setae basally as in Fig. 
140, setae very fine, costal cell almost naked, 
linea calva open posteriorly; postmarginal vein 
nearly absent. Relative measurements (holo- 
type): forewing length 80, forewing width 36; 
hindwing length 59; hindwing width 12. 

Gaster about as long as thorax; ovipositor not 
exserted, similar to Fig. 335; hypopygium similar 
to Fig. 298. Relative measurements: ovipositor 
length 99, gonostylus 19 [mid tibia 112]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. India. 

Material examined. Type material. Holotype 
O, INDIA: Delhi, 11-18.V.1985 (J. LaSalle). 
Paratype, 1$, same data as holotype. Material in 
BMNH. 

Comments. See the comments under Ooencyr- 
tus elissa (above). 

Ooencyrtus erebus sp.n. 

(Figs 141-147) 

Diagnosis. Female: body generally dark brown 
or blackish, frontovertex weakly metallic pur- 
plish, mesoscutum bluish and scutellum contrast- 
ing coppery purple margined green at apex and 
sides; antennae generally testaceous; legs yellow, 
but mid and hind coxae black; ovipositor sheaths 
brown; mandibles (Fig. 143) with two teeth and a 
truncation; frontovertex about one-quarter head 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



45 



width; flagellum (Figs 141, 142) filiform, gener- 
ally F1-F5 longer than broad, F6 subquadrate; 
clava hardly broader than F6, its sutures parallel, 
sensory area small, at extreme apex only; scutel- 
lum (Fig. 146) with shallow sculpture as mesoscu- 
tum or shallower; extreme apex and sides smooth 
and shiny, delimited by a shallow carina; apex of 
scutellum rounded, sometimes slightly produced 
(Fig. 146); forewing (Figs 147) with a distinct 
naked basal area; ovipositor hardly exserted. 
Male: generally similar to female but antennae 
yellow, face shining green; antennae with all 
funicle segments except Fl at least 2 times as 
long as broad, Fl shorter; setae on flagellum at 
least 2 times as long as diameter of segments; 
frontovertex a little more than one-third head 
width. 

Female. Length about 0.80-1.15 mm (holotype 
0.92 mm). 

Head blackish, weakly shining purplish on 
frontovertex; face greenish but with interanten- 
nal prominence quite metallic purple; radicle 
dark brown, rest of antenna testaceous or testa- 
ceous yellow occasionally with clava conspicu- 
ously darker; pronotum and axillae purplish; 
mesoscutum shining green or blue-green; scutel- 
lum shining coppery purple with margins and 
sides metallic green; mesopleuron dark purplish 
brown; propodeum black; legs completely yellow 
but with mid and hind coxae dark brown; wings 
hyaline, venation testaceous, marginal vein 
brown; gaster black with weak green and purple 
sheen; ovipositor sheaths brown. 

Head with shallow, regular, polygonally 
reticulate sculpture on frontovertex; lower face 
and genae with shallower, longitudinally elon- 
gate sculpture; ocelli forming an approximately 
equilateral triangle; posterior ocelli separated 
from occipital margin by a little more than their 
own lengths and conspicuously separated from 
eye margins; frontovertex about one-quarter 
head width; eyes not conspicuously hairy; occipi- 
tal margin sharp; antennae inserted well below 
the ventral eye margins; toruli separated from 
mouth margin by a little less than their own 
lengths; antennae (Figs 141, 142) with pedicel 
plus flagellum about 1.1 times as long as head 
width; Fl-5 conspicuously longer than broad, F6 
subquadrate; clava hardly broader than F6 and 
nearly as long as F3-6 together, with sutures 
parallel and sensory area small and limited to 
extreme apex only; mandibles (Fig. 143) with 
two teeth and a truncation. Relative measure- 
ments (holotype): head width 35, head height 31, 
minimum frontovertex width 9, POL 4, OPL 3, 
OOL 1, eye length 22, eye width 18, scape length 



16, scape width 3, malar space 12; other propor- 
tions of antennae as in Fig. 142. 

Mesoscutum with shallow imbricate sculpture; 
scutellum (Fig. 146) with similar sculpture 
basally but becoming shallower towards sides 
and apex, laterally more elongate; extreme sides 
and apex of scutellum smooth and polished, 
clearly separated by a shallow carina or impres- 
sion, apex more or less rounded; forewing with 
venation and distribution of setae basally as in 
Figs 144 and 147, with a distinct naked basal area 
with only a few setae posteriorly on ventral 
surface of basal cell; linea calva closed posteri- 
orly or nearly so (Figs 144, 147). Relative mea- 
surements (paratype): forewing length 87, 
forewing width 35; hindwing length 60; hindwing 
width 12. 

Gaster with ovipositor hidden or hardly 
exserted; ovipositor as in Fig. 145; hypopygium 
similar to Fig. 149. Relative measurements 
(paratype): ovipositor length 102, gonostylus 24 
[mid tibia 75]. 

Male. Length about 0.7 mm. Generally similar 
to female, but for coloration and structure of 
antennae, wider frontovertex and structure of 
genitalia. Antennae completely yellow; pedicel 
plus flagellum 1.75 times as long as head width, 
funicle segments subequal in length and about 2 
times as long as broad except Fl which is only 
slightly longer than broad; flagellum clothed in 
setae at least 2 times as long as diameter of 
segments; clava a little shorter than F5-6 com- 
bined; frontovertex about two-fifths head width; 
ocelli forming a right angle; posterior ocelli sepa- 
rated from occipital margin by a little more than 
their own lengths. 

Variation. In the female the coloration of the 
antennae, head and thorax varies as outlined 
above; the antennae sometimes have Fl-2 quad- 
rate and smaller than F3-6 (Fig. 141) and the 
linea calva is sometimes nearly closed posteriorly 
(Fig. 144). 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, iv.1985 (J.S. Noyes). 
Paratypes, 4$, 2d\ same data as holotype; 15$, 
Sulawesi Tengah, Nr Morowali, Ranu River 
Area, Lowland rain forest, B.M. 1980-280, 
hi. 1980 (M.J.D. Bredell). Holotype in BMNH, 
paratypes in BMNH, IZAS, MZB. 

Comments. Females of O. erebus can be sepa- 



46 



D. -W.HUANG AND J.S. NOYES 



rated from other species of the genus by the legs 
being completely yellow except for the mid and 
hind coxae, and the strongly contrasting metallic 
blue or green mesoscutum with the coppery 
purple scutellum. 

Ooencyrtus endymion sp.n. 

(Figs 148-153) 

Diagnosis. Female: body generally dark 
brown; head and dorsum of thorax metallic 
green; fore and mid legs almost completely yel- 
low, hind coxae and femora brown, mid and hind 
tibiae each with a faint brown subbasal ring; 
ovipositor with gonostylus dark brown; scutellum 
with similar sculpture to mesoscutum, its apex 
slightly pointed; antennae with radicle, scape and 
pedicel dark brown, flagellum testaceous; flagel- 
lum more or less filiform; all funicle segments 
longer than broad; clava hardly not broader than 
F6, sutures subparallel and sensory part at 
extreme apex only; forewing (Fig. 153) with only 
a very small naked basal area; linea calva closed 
posteriorly; mandibles (Fig. 150) with two teeth 
and a truncation; frontovertex about one-quarter 
head width; ovipositor not exserted. Male: 
unknown. 

Female. Length about 1.11-1.13 mm (holotype 
1.11mm). 

Head metallic green sometimes slightly bluish 
and ocellar area slightly coppery, interantennal 
prominence purple, but with a narrow blue line 
between toruli; antennae with radicle, scape and 
pedicel brown; flagellum testaceous; dorsum of 
thorax metallic green, occasionally scutellum 
slightly bluish; mesopleuron and propodeum 
dark purple-brown; legs mostly yellow, but hind 
coxae dark brown, hind femora brown with api- 
ces yellow, mid and hind tibia each with a 
narrow, faint, subbasal brown ring; wings hya- 
line, venation testaceous; gaster dark purple- 
brown with a weak green sheen; ovipositor 
sheaths brown. 

Head with fairly regular, shallow, raised 
polygonal sculpture on frontovertex; lower face 
and genae with shallow longitudinally elongate 
sculpture; ocelli forming an angle of about 
70-80°; posterior ocelli separated from occipital 
margin by about 1.5 times their own lengths and 
conspicuously separated from eye margins; fron- 
tovertex about one-quarter head width; eyes 
fairly conspicuously hairy; occipital margin 
hardly rounded; antennae inserted well below 
the ventral eye margins; toruli separated from 
mouth margins by about their own lengths or a 
little less; clypeus very slightly convex medially; 



antennae (Fig. 148) with pedicel plus flagellum as 
long as head width; Fl-5 conspicuously longer 
than broad, F6 slightly longer than broad; clava 
hardly broader than F6 and a little longer than 
F4-6 together, with sutures parallel, sensory area 
slightly enlarged but at extreme apex only; man- 
dibles (Fig. 150) normally with two teeth and a 
truncation, but in at least one specimen they are 
broadly truncate, without teeth (Fig. 151). Rela- 
tive measurements (holotype): head width 52, 
head height 43, minimum frontovertex width 
13.5, POL 8, OPL 3.5, OOL 1, eye length 30, 
eye width 26, scape length 22, scape width 5, 
malar space 17; other proportions of antennae as 
in Fig. 148. 

Mesoscutum with shallow, regular imbricate- 
reticulate sculpture; scutellum with similar, but 
slightly deeper sculpture basally, this becoming 
gradually more shallow towards apex and more 
elongate and shallower towards sides; apex mod- 
erately pointed; forewing with venation and dis- 
tribution of setae basally as in Fig. 153, with a 
distinct, enclosed basal naked area; linea calva 
closed posteriorly. Relative measurements (holo- 
type): forewing length 135, forewing width 61; 
hindwing length 98; hindwing width 26. 

Gaster shorter than thorax; ovipositor (Fig. 
152) not exserted, or hardly so, with gonostylus 
about one-fifth as long as ovipositor, second 
valvifer proximally strongly curved; hypopygium 
as in Fig. 149. Relative measurements 
(paratype): ovipositor length 59, gonostylus 11 
[mid tibia 51]. 

Male. Unknown. 

Variation. There is some variation in colora- 
tion of the head and thorax as described above 
with the head and scutellum having a distinct 
bluish lustre contrasting with the green mesoscu- 
tum. At least one of specimen from Sichuan has 
edentate mandibles, but this may be because the 
mandibles are worn. These specimens are also 
conspicuously smaller than those from Yunnan 
and consequently the antennal segments are rela- 
tively shorter. 

Hosts. Recorded from the eggs of Dendrolimus 
sp., Dendrolimus kikuchii (Lepidoptera; Lasio- 
campidae) and an unidentified sphingid. 

Distribution. P.R. China. 

Material examined. Type material. Holotype 
$, P.R. CHINA: Yunnan, ex eggs of Dendroli- 
mus sp., 83-1285, 28.viii.1981 (Hou Taoqian). 
Paratypes, P.R.CHINA: 1$, same data as holo- 
type; 5$, Yunnan, Gejiu, ex eggs of Dendroli- 
mus kikuchii Matsumura, 10.viii.1980 (Li 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



41 



Guoxiu); 39, Yunnan, Honghe, ex. eggs of a 
sphingid (Li Guoxiu); 3$, Sichuan, Huili, Hua- 
ngxian, 2.vi.l961 (Dingxi Liao). Holotype in 
IZAS, paratypes in IZAS, BMNH, 

Comments. Ooencyrtus endymion is very close 
to pinicolus (Matsumura) recorded from the 
same host genus in the Palaearctic region. 
Females of O. pinicolus are generally much 
darker than endymion with all coxae dark brown 
and the rest of the legs almost completely dark 
brown. Most notably the frontovertex of pinico- 
lus is nearly one-third head width and the linea 
calva is open, whereas in endymion the linea 
calva is distinctly closed and the frontovertex is 
about one-quarter head width. O. endymion is 
also very similar to O. fecundus Ferriere & 
Voegele from North Africa, both species being 
of similar general habitus and coloration, espe- 
cially the legs and dorsum of thorax. O. fecundus 
has also been recorded from the eggs of Lasio- 
campidae and Sphingidae (Trjapitzin, 1989). 
However, in fecundus the hind femora are com- 
pletely yellow, the postmarginal vein is very 
short and almost absent and the linea clava is 
open, whereas in endymion the hind femora are 
mostly dark brown, the postmarginal vein is 
about two-thirds as long as the stigmal and the 
linea calva is closed. 

Ooencyrtus hymen sp.n. 

(Figs 154-158) 

Diagnosis. Female: head largely metallic 
green, lower parts of face orange; thorax orange 
mixed with metallic green dorsally; antennae and 
legs orange; gaster entirely dark brown with 
weak metallic reflections; forewings hyaline; ovi- 
positor sheaths pale orange; mandible with two 
teeth and a narrow emarginate truncation (Fig. 
155); antennae (Fig. 154) with funicle segments 
at most slightly longer than broad, with clava 
slightly enlarged about as long as F3-6 together, 
its apex with a very small oblique apical trunca- 
tion and enlarged sensory area; frontovertex 
extremely narrow, narrowest anterior to ocelli 
and about one-seventeenth head width; posterior 
ocelli nearly touching, forming a very acute 
angle; median ocellus almost touching eyes; eyes 
conspicuously hairy; forewing (Fig. 156) without 
a conspicuous basal naked area; linea calva 
closed or nearly so; scutellum with similar sculp- 
ture to mesoscutum, only posterior vertical mar- 
gin smooth and shiny; ovipositor very slightly 
exserted. Male: unknown. 



Head metallic green on frontovertex, upper 
scrobal area and temples, lower face including 
interantennal prominence and genae orange; 
antennae including radicle orange; thorax 
orange, but distinctly metallic green medially on 
dorsum, and on scutellum; tegulae orange; legs, 
including coxae, pale orange; forewing hyaline, 
veins testaceous; propodeum orange; gaster com- 
pletely dark brown with a metallic coppery and 
green sheen; ovipositor sheath pale orange. 

Head with very fine, polygonally reticulate 
sculpture on frontovertex; sculpture on genae 
longitudinally elongate; ocelli forming a very 
acute angle of about 15°, the posterior ocelli 
almost touching and separated from occipital 
margin by nearly twice their own lengths and 
touching eye margins; median ocellus nearly 
touching eyes; frontovertex, at narrowest point 
about mid way between anterior ocellus and top 
of scrobes, one-seventeenth head width; eyes 
conspicuously hairy; occipital margin acute, 
hardly rounded; head in front view (Fig. 158) 
nearly as broad as high; antennae inserted far 
below the ventral eye margins, toruli separated 
from mouth margin by their own lengths; anten- 
nae (Fig. 154) with pedicel plus flagellum about 
as long as head width; pedicel distinctly longer 
than Fl-2 together; Fl-5 subquadrate, F6 trans- 
verse, flagellum clavate; clava about as long as 
F3-6 together, the sutures parallel but apex with 
a small, slightly oblique truncation, the sensory 
part slightly enlarged; mandibles almost triden- 
tate, with two teeth and a short, slightly emargin- 
ate truncation. Relative measurements 
(holotype): head width 53, head height 52, mini- 
mum frontovertex width 3, POL 2, OPL 5, OOL 
0, eye length 41, eye width 32, malar space 18, 
scape length 22, scape width 4.5; other propor- 
tions of antenna as in Fig. 154. 

Scutellum with very similar fine imbricate- 
reticulate sculpture to mesoscutum, only its pos- 
terior vertical margin smooth; forewing hyaline 
with venation and distribution of setae basally as 
in Fig. 156, without a conspicuous basal naked 
area and linea calva closed or nearly so. Relative 
measurements (holotype): forewing length 120, 
forewing width 50; hindwing length 74; hindwing 
width 19. 

Gaster about as long as thorax; ovipositor 
(Fig. 157) very slightly exserted; hypopygium 
similar to Fig. 218. Relative measurements 
(paratype): ovipositor length 53; gonostylus 14 
[mid tibia 45]. 

Male. Unknown. 



Female. Length 1.35 mm. (holotype 1.35 mm). Variation. The extent of the metallic green 



48 



coloration of the mesoscutum is variable, it may 
be almost absent, limited to an inconspicuous 
area medially or it may be more extensive and 
covering almost the whole of the mesoscutum 
with only the anterior and outer margin orange 
as in the holotype. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 300m, BMNH Canopy Fog 
#4, 8.ii. 1985. Paratypes, 1$, same data as holo- 
type except BMNH Canopy Fog #3 and without 
elevation; 1$, 450m, BMNH Canopy Fog. #5, 
2.ii.l985, other data as holotype. Material in 
BMNH. 

COMMENTS. Ooencyrtus hymen is very close to 
belus, females of both species having the thorax 
largely yellow or orange, legs completely yellow 
or orange, wings hyaline with linea calva closed 
or nearly so, narrow frontovertex, conspicuously 
hairy eyes and more or less uniformly sculptured 
scutellum. The two species can be separated by 
the relative size of the sensory area of the clava 
and by the coloration of the dorsum of the thorax 
(see key). 

Ooencyrtus belus sp.n. 

(Figs 159-165) 

Diagnosis. Female: head and dorsum of thorax 
almost entirely metallic green or blue-green, 
sides of thorax, antennae and legs orange; gaster 
completely dark brown, ovipositor sheaths pale 
orange; forewing hyaline; antennae (Fig. 160) 
filiform; funicle segments at most slightly longer 
than broad; pedicel plus flagellum as long as 
head width; clava a little longer than F3-6 
together, rounded at apex and without a con- 
spicuous sensory area; frontovertex about one- 
seventeenth head width, narrowest below 
anterior ocellus; mandible with two teeth and 
short truncation (similar to Fig. 155); posterior 
ocelli almost touching, median ocellus almost 
touching eyes (Fig. 159); eyes conspicuously 
hairy; forewing (Fig. 161) without a conspicuous 
naked basal area; linea calva nearly closed; 
scutellum with sculpture similar to that on mesos- 
cutum, only posterior vertical margin smooth; 
ovipositor slightly exserted. Male: generally very 
similar to female but scape expanded in proximal 
half, flagellum white, funicle segments all trans- 
verse and clothed with setae that are not longer 
than diameter of segments and clava enlarged 



D. -W.HUANG AND J.S. NOYES 
with an oblique apical truncation (Fig. 164). 

Female. Length (including ovipositor) 
0.90-0.95 mm (holotype 0.95 mm). 

Head metallic blue-green, mouth margin 
orange, interantennal prominence purple-blue; 
antennae including radicle orange; sides of tho- 
rax and legs orange, dorsum metallic green; 
tegulae orange; propodeum brown; gaster 
entirely with a coppery purple and green sheen; 
forewing hyaline, veins testaceous; ovipositor 
sheaths yellowish. 

Head with fine polygonally reticulate sculpture 
on frontovertex; sculpture on genae longitudi- 
nally elongate; ocelli forming a very acute angle 
of about 15°, posterior ocelli nearly touching and 
separated from occipital margin by nearly twice 
their own lengths and touching eye margins; 
median ocellus nearly touching eyes (Fig. 159); 
frontovertex about one-seventeenth head width; 
eyes conspicuously hairy; occipital margin acute, 
hardly rounded; head in front view (Fig. 159) 
only slightly broader than high; antennae 
inserted far below ventral eye margin, toruli 
separated from mouth margin by less than their 
own lengths; antennae (Fig. 160) with pedicel 
plus flagellum as long as head width; pedicel 
distinctly longer than Fl plus F2 and slightly 
broader than Fl; Fl^ subquadrate, F5 and F6 
transverse, flagellum filiform; clava a little longer 
than F3-6 together, rounded at apex and without 
a conspicuous apical sensory area; mandible 
appearing almost tridentate with two teeth and a 
very short truncation (similar to Fig. 155). Rela- 
tive measurements (holotype): head width 41, 
head height 39, minimum frontovertex width 2.5, 
POL 2, OPL 4, OOL 0, eye length 31, eye width 
26, malar space 13, scape length 16, scape width 
4; other proportions of antenna as in Fig. 160. 

Scutellum with similar, fine, imbricate- 
reticulate sculpture as mesoscutum, only poste- 
rior vertical margin smooth; forewing (Fig. 161) 
hyaline, without a conspicuous basal naked area 
and linea calva posteriorly nearly closed. Rela- 
tive measurements (holotype): forewing length 
92 forewing width 37; hindwing length 58; hind- 
wing width 14. 

Gaster a little shorter than thorax; ovipositor 
(Fig. 162) slightly exserted; hypopygium as in 
Fig. 163. Relative measurements: ovipositor 
length 60, gonostylus 16 [mid tibia 35]. 

Male. Length 0.72 mm. Extremely similar to 
female but for antennae and genitalia. Antennae 
(Fig. 164) with scape and pedicel yellow, flagel- 
lum white; scape expanded in proximal half; 
pedicel longer than proximal funicle segments 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



49 



together; at most F1-F3 subquadrate, F4-F6 
transverse; clava enlarged with an oblique apical 
truncation, and with very distinct sensory tubules 
on the truncate part; genitalia as in Fig. 165, with 
a single tooth on each digitus. 

Variation. Very little in material available. 

Hosts. Recorded below from mealybugs 
(Homoptera: Pseudococcidae) on coconut, but 
this is almost certainly erroneous. The specimens 
probably originated from insect eggs mixed in 
with the mealybugs. 

Distribution. Philippines 

Material examined. Type material. Holotype 
9, PHILIPPINES: Christensen, Plant. 
Hagonoy, Davao del Sur, ex mealybug on coco- 
nut, CLE. A16554, i.1984. Paratypes, 3$, 2d\ 
same data as holotype. Material in BMNH. 

Comments. Ooencyrtus belus is close to hymen 
(see comments under hymen). 

Ooencyrtus bacchus sp.n. 

(Figs 166-168) 

Diagnosis. Female: generally blackish, dark 
metallic green on head, greenish and brassy on 
mesoscutum and bluish on scutellum; a thin, 
faint purplish u-shaped line connects the toruli 
ventrally; legs, including coxae yellow; scape, 
pedicel and clava yellow, flagellum brown; gaster 
blackish with a slight green and brassy sheen; 
ovipositor sheaths yellow; mandibles with one 
tooth and a broad inner, minutely denticulate 
truncation; frontovertex about one-fourth head 
width; funicle segments at most quadrate, slightly 
wider distally, F5-6 conspicuously transverse; 
clava distinctly broader than funicle segments, 
without an enlarged apical sensory area; clypeus 
very slightly convex medially; scutellum with 
similar sculpture to mesoscutum in anterior one- 
third, posterior two-thirds smooth and shiny; 
forewing (Fig. 168) without a distinct naked basal 
area; linea calva open posteriorly; gaster shorter 
than thorax; ovipositor not exserted. 

Female. Length 0.70-0.75 mm (holotype 0.75 
mm). 

Head metallic green, coppery purple in ocellar 
area and brassy on genae; lower margins of toruli 
connected by a very thin, inconspicuous metallic 
purplish line across the lower part of the interan- 
tennal prominence; antennae with radicle black, 
scape and pedicel orange, funicle dark brown 
and clava yellow; pronotum, dark purple-brown; 
mesoscutum weakly metallic green, bordered 



purplish, axillae black with weak green tinge; 
scutellum anteriorly blackish and in posterior 
two-thirds metallic blue; mesopleuron and pro- 
podeum dark brown; all legs, including coxae, 
yellow; wings hyaline, venation testaceous; 
gaster dark purple-brown with slight green, 
purple or brassy sheen; ovipositor sheaths yel- 
low. 

Head with shallow, regular, polygonally 
reticulate sculpture on frontovertex; sculpture on 
lower face and genae shallower and longitudi- 
nally elongate; ocelli forming an equilateral tri- 
angle; posterior ocelli separated from occipital 
margin by about twice their own lengths and 
nearly touching eye margins; frontovertex about 
one-quarter head width; eyes not conspicuously 
hairy; occipital margin slightly rounded; anten- 
nae inserted just below ventral eye margins; 
toruli separated from mouth margin by about 
their own lengths; clypeus slightly convex medi- 
ally; mandibles with a sharp outer tooth and a 
broad inner truncation bearing some mini-teeth 
on inner half; antennae (Fig. 167) with pedicel 
plus flagellum about as long as head width; scape 
at least 4 times as long as broad; funicle segments 
quadrate or transverse; clava distinctly broader 
than F6, as long as F3-6 together, with sensory 
area small and at apex only. Relative measure- 
ments (holotype): head width 36, head height 31, 
minimum frontovertex width 9, POL 4, OPL 3.5, 
OOL 0.3, eye length 22, eye width 17, scape 
length 15, scape width 3.5, malar space 13; other 
proportions of antennae as in Fig. 167. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum with polygonally reticulate 
sculpture anteriorly, not deeper than mesoscu- 
tum, gradually becoming shallower posteriorly, 
posterior two-thirds smooth, apex rounded; 
forewing with venation and distribution of setae 
basally as in Fig. 168, without a distinct naked 
basal area; linea calva open posteriorly. Relative 
measurements (paratype): forewing length 84, 
forewing width 35; hindwing length 59; hindwing 
width 12. 

Gaster shorter than thorax, last tergite some- 
what emarginate posteriorly; ovipositor (similar 
to Fig. 335) hidden; hypopygium as in Fig. 166. 
Relative measurements (paratype): ovipositor 
length 76, gonostylus 13 [mid tibia 79]. 

Male. Unknown. 

Variation. Very little in material available. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 



50 



D. -W.HUANG AND J.S. NOYES 



$, INDONESIA: Rakata Kecil, Krakatau Cent. 
Expdn, 20. ix. 1984 (S.G. Compton). Paratypes, 
35 (one on slide), same data as holotype. Holo- 
type in BMNH, paratypes in BMNH and MZB. 

Comments. Ooencyrtus bacchus probably 
belongs to the same group of species as ceres, iris 
and elissa, all four species having the clypeus 
slightly convex medially, a very thin inconspicu- 
ous purple line connecting the lower margins of 
the toruli and very similar mandibular structure 
with the truncate part minutely denticulate. Of 
these species it is most similar to elissa in general 
habitus and coloration but can be separated from 
this species by the relatively shorter funicle seg- 
ments and completely yellow legs. O. elissa has 
F3-6 conspicuously longer than broad and the 
hind coxae dark brown and the hind femur with a 
dark brown dorsal stripe. O. bacchus may also be 
close to pacificus, both species having all coxae 
yellow, clava distinctly paler than the funicle, 
clypeal margin medially produced, mandibles 
with a minutely denticulate truncation, posterior 
two-thirds of the scutellum smooth; forewing 
without conspicuous naked basal area and linea 
calva open posteriorly. In addition to the charac- 
ters given in the key O. bacchus can be separated 
from pacificus by having the hind femora com- 
pletely yellow, the clava distinctly broader than 
F6 and at least as long as F4-6 together, and 
different structure of the hypopygium (compare 
Figs 166 and 171). In pacificus the hind femora 
are apically infuscate and the clava hardly 
broader than F6 and not as long as F3-6 
together. 

Ooencyrtus pacificus Waterston 

(Figs 169-174) 

Ooencyrtus pacificus Waterston, 1915: 307-310. 
LECTOTYPE $, (here designated) Fiji 
(BMNH, examined). 

Diagnosis. Female: length about 0.8-0.9 mm; 
body generally dark brown or blackish, head 
metallic green, genae somewhat brassy, interan- 
tennal prominence purple; mesoscutum metallic 
blue-green, posterior two-thirds of scutellum 
metallic green; antennae (Fig. 169) blackish with 
clava and tip of F6 contrasting pale yellow; legs 
completely yellow but with hind-femora a little 
infuscate apically; forewing almost hyaline but 
with an inconspicuous transverse infuscate band 
from apex of venation (Fig. 170); mandible (Fig. 
173) with one tooth and a broad, minutely den- 
ticulate truncation; scape about 3 times as long as 
broad, pedicel slightly longer than Fl-2 together; 



Fl-3 smaller than F4-6; frontovertex about one- 
quarter head width; scutellum anteriorly with 
shallower sculpture than mesoscutum, posterior 
two-thirds or so smooth and shiny; forewing (Fig. 
170) without a conspicuous naked basal; linea 
calva open; postmarginal vein about one-third as 
long as stigmal; ovipositor (Fig. 171) not 
exserted; hypopygium similar to Fig. 36. Male: 
length 0.6-0.8 mm; generally similar to female 
but flagellum unicolorous testaceous-brown; 
hind-femora dark brown; all funicle segments at 
least twice as long as broad and clothed in setae 
2-3 times as long as diameter of segments (Fig. 
174); aedeagus about 0.6 times as long as mid 
tibia. 

Hosts. Reared from eggs of the bean bug (Brac- 
hyplatys pacificus) (Waterston, 1915), and 
recorded below from an unidentified pest of 
pigeon pea. 

Distribution. Fiji. 

Material examined. Type material. Lecto- 
type 9- FIJI: Rarawai, Viti Levu, bred from eggs 
of the Bean Bug (Brachyplatys pacificus Dall.) 
(R. Veitch), B.M. TYPE HYM. 5. 1,076. Para- 
lectotypes, 1$, lcf, same data as lectotype, but 
no type number (BMNH). 

Other material. FIJI: 89, 3o\ Viti Levu, ex 
pests of pigeon pea, Lab. bred, HE 22057, 
26.ix.1991. Material in BMNH. 

Comments. A very detailed description of this 
species is provided by Waterston (1915). See also 
comments under Ooencyrtus bacchus above. 

Ooencyrtus larvarum (Girault) 

(Figs 175-180) 

Paracopidosomopsis larvarum Girault, 1919b: 
58. LECTOTYPE 9 (here designated), Indo- 
nesia (QMB, examined). 

Ooencyrtus larvarum (Girault); Noyes & Hayat, 
1984: 309. 

Diagnosis. Female (length 0.71 mm-0.80 mm): 
body generally dark brown or blackish; head, 
mesoscutum and basal two-thirds of scutellum 
with a dull metallic green lustre; apical third of 
scutellum smooth and bright metallic green; 
scape and pedicel yellowish, flagellum 
testaceous-yellow or brownish; tegulae basally 
yellow; legs, including coxae, yellow; gaster dark 
brown; antennae (Fig. 175) with Fl transverse 
and much smaller than F2-6 which are all trans- 
verse; clava about as long as F4-6 together and 
with a very slight oblique apical truncation and 
sensory enlarged somewhat enlarged; mandibles 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



51 



(Fig. 176) with two long sharp teeth and a small, 
inner third tooth; frontovertex about one-third 
head width; gena, about two-thirds eye length, 
mouth opening less than two-fifths head width 
with clypeus medially distinctly convex (Fig. 
178); ovipositor (Fig. 179) with basal part of 
second valvifer characteristically broadly curved, 
but short, gonostylus one-quarter ovipositor 
length; hypopygium (Fig. 177) short. Male 
(0.63-0.71 mm): very similar to female but for 
antennae and genitalia. Antennae (Fig. 180) with 
clava yellowish and slightly paler than rest of 
flagellum; all funicle segments subquadrate, but 
Fl clearly smaller than other segments which are 
subequal; flagellum clothed in setae which are 
not longer than diameter of segments; aedeagus 
about two-thirds as long as mid tibia. 

Hosts. The type series was reared from larva of 
Delias sp. (Girault, 1919b), but material 
recorded below was reared from eggs of Eurema 
sp. (both Lepidoptera: Pieridae). 

Distribution. Thailand, Indonesia. 

Material examined. Type material. Lecto- 
type 9- 'Paracopidosomopsis larvarum Gir. 9 
Types' in Girault's handwriting, QUEENS- 
LAND MUSEUM [green label], no other data 
but according to Girault ( 1919b) the material was 
reared from a larva of Delias sp. from Salatiga, 
Java in INDONESIA. Paralectotypes, 2$, 79. 
same data as lectotype, but mounted separately 
(19' 2o" remounted on slides). Material in 
QMB. 

Non-type material. THAILAND: 29, 2d\ 
Chedi Mae Khrua, ex eggs of Eurema sp., 26. i/ 
5.H.1990 (R. Harberd). Material in BMNH. 

Comments. Girault (1919b) mentions only 
males, but mounted on the same card as the male 
syntypes are three females and therefore we have 
no hesitation in selecting the best of these as 
lectotype. Superficially males and females of this 
species are very similar, especially if poorly 
mounted as are the syntypes. 

This species has very characteristic mandibles 
(see Fig. 176) and antennae (Fig. 175) and both 
sexes should prove to be easy to separate from 
other species of the genus on these characters 
alone. The head shape is also very characteristic 
being similar to species of Arrhenophagus with 
elongate genae which taper inwards towards a 
relatively narrow mouth and a distinctly convex 
clypeal margin. 



Ooencyrtus pallidipes (Ashmead) 

(Figs 181-186) 

Aphidencyrtus pallidipes Ashmead, 1904a: 15. 
LECTOTYPE 9 (here designated), Philip- 
pines (USNM, examined). 

Ooencyrtus erionotae Ferriere, 1931: 284. LEC- 
TOTYPE 9 (here designated), Malaysia 
(BMNH, examined). Syn. n. 

Ooencyrtus pallidipes (Ashmead); Noyes & 
Hayat, 1984: 309. 

Diagnosis. Body generally dark brown or 
blackish; head largely metallic green, purplish in 
ocellar area and on interantennal prominence; 
mesoscutum dull bluish green; scutellum dull 
green anteriorly with slightly coppery tinge, shin- 
ing metallic green in posterior one-quarter or so; 
flagellum testaceous-yellow; all legs, including 
coxae, yellow; mandibles with one tooth and a 
broad truncation; scutellum anteriorly with 
punctate-reticulate sculpture and with posterior 
one-quarter smooth (similar to Fig. 316); forew- 
ing (Figs 182, 184) with large naked basal area; 
basal cell with a small patch of ventral setae 
posteriorly; linea calva open; postmarginal vein 
from one-third to two-thirds length of stigmal. 
Female (length 0.82-1.25 mm): gaster blackish 
but sometimes with small whitish marks laterally 
near petiole dorsally; ovipositor sheaths yellow; 
antennae (Fig. 181) filiform, Fl-5 distinctly 
longer than broad; F6 slightly transverse to 
slightly longer than broad; frontovertex at most 
1/5 head width (Fig. 183); ovipositor very slightly 
exserted and similar to Fig. 122, 1.2 times as long 
as mid-tibia, gonostylus about one-sixth oviposi- 
tor length; hypopygium as in Figs 185, 186. Male 
(length 0.75-0.95 mm): as female but antennae 
whitish or pale yellow, Fl smaller than F2-6 but 
all funicle segments longer than broad and 
clothed in setae a little longer than diameter of 
segments; frontovertex nearly one-third head 
width. 

Hosts. A parasitoid of the eggs of Erionota 
thrax (Lepidoptera: Hesperiidae), but also 
recorded from the eggs of Caligo memnon (Lepi- 
doptera: Nymphalidae) a Neotropical species 
being reared in Thailand for commercial pur- 
poses (see below). The original record from an 
undetermined aphid host (Ashmead, 1904a) is 
undoubtedly erroneous. The record of this spe- 
cies, as O erionotae, from the eggs of Agrilus 
sexsignatus (Coleoptera: Buprestidae) (Braza, 
1988, 1989) is a misidentification of Orianos 
brazai Noyes (see Noyes, 1990). 



52 



D. -W.HUANG AND J.S. NOYES 



Use in biocontrol. The species was succesfully 
introduced into Mauritius in 1971-1972 (Water- 
house & Norris, 1989) and Hawaii in 1973 (Mau 
et al., 1980) for the control of Erionota thrax 
(Lepidoptera: Hesperiidae). 

Distribution. Mauritius (introduced), Nepal, 
Thailand, P. R. China, Philippines, Malaysia, 
Indonesia, Papua N. Guinea, Guam, Hawaii 
(introduced). 

Material examined. Type material. Aphiden- 
cyrtus pallidipes Ashmead: Lectotype 9 - PHILIP- 
PINES: 'Manila PI, W.A. Stanton Collector, 
Aphidencyrtus pallidipes Ash. cf $' in Ashmead's 
handwriting, '$ Type No.7701 U.S.N.M.' 
(USNM). Paralectotype, 1$, same data as lecto- 
type (BMNH). Ooencyrtus erionotae Ferriere: Lec- 
totype 9, MALAYSIA: 'Type, Malaya, Sungei 
Tua, 31.vii.28, ex. ova of Erionota thrax, G.H. 
Corbett, Pres. by Imp. Inst. Ent. Brit. Mus. 
1931-367, Ooencyrtus erionotae sp.n. 9 type, Ch. 
Ferriere det., B.M.TYPE HYM. 5. 1,071' 
(BMNH). Paralectotypes, MALAYSIA: 19, 39, 
same data as lectotype but with 'cotype' label; 199 > 
3d", Kuala Lumpur, 13. i. 1921, Ulu Gombak, ex. 
ova of indet. on wild plantain leaf, 27. i. 1921 (Dr, 
W.A. Lamborn) 'Ooencyrtus erionotae, sp.n. Ch. 
Ferriere det.' (BMNH). 

Non-type material. NEPAL: 39, Id, Ram- 
pur, ex. eggs of banana skipper, 2.vi.l981; 
THAILAND: 79, Id", Chedi Mae Khrua , ex 
eggs of Caligo memnon (N.), 23.i/ll.ii.l990 (Ray 
Harberd); 39, Chedi Mae Khrua, ex eggs of 
unknown species, 7.1990 (Ray Harberd); P. R. 
CHINA: 109, 2d, Fujian, Fuzhou, ex. eggs of 
Erionota thrax on banana, 5.vii.l986; 259, 3d", 
Hainan, Xinglong, ex. eggs of butterfly, 
5.iv.l964 (D.-x. Liao); INDONESIA: 1$, East 
Sumatra, Asahab 50-60m., from gambir pests, 
1934-1936 (F. Schneider); 1$, Java, Siloewok, 
Sawangan Est., ex eggs of Hadart thrax, 
18.viii.1938 (Ir Duyvendijk); PAPUA NEW 
GUINEA: 39, Id, Morobe Prov., Markham 
Velley, Kaiapit, ex eggs Erionota thrax on 
banana, 15. v. 1987 (P. Merrett); 139, lCf, Lae, 
ex Erionota thrax, 13.vi.1987 (P. Merrett); 39, 
Lae, ex eggs of Erionota thrax, 8.viii.88; 59, 
West Highland Province, Mt. Hagen, ex. eggs of 
banana skipper, CIE A20946, 27. vi. 1989 (T. 
Solulu); 169 , lcf, Morobe Prov., ex. eggs of 
Erionota thrax, CIE A19626, 15.V.1987 (P. J. 
Merrett). In BMNH. 

Comments. Ooencyrtus pallidipes is generally 
similar to pilosus (see comments under pilosus 
below). 



Ooencyrtus pilosus sp.n. 

(Figs 187-189) 

Diagnosis. Female: body generally dark brown 
or blackish; head metallic green with a coppery 
purple sheen; mesoscutum with a weak blue- 
green lustre; scutellum green; antennae yellow- 
ish; legs, including coxae, yellow; ovipositor 
sheaths dark brown; antennae (Fig. 187) with 
flagellum filiform; pedicel broader than Fl and 
as long as Fl-2 combined; all funicle segments 
longer than broad, Fl-2 smaller and more slen- 
der than F3-6; clava slender and apically 
rounded; frontovertex about one-quarter to one- 
sixth head width; ocelli forming an acute angle; 
mandibles with one tooth and a broad, minutely 
denticulate truncation; forewing (Fig. 189) with 
marginal vein distinctly longer than broad, basal 
cell densely hairy without a conspicuous naked 
area basally, linea calva closed posteriorly; 
scutellum slightly convex, with posterior one- 
fifth or so smooth and shiny, anteriorly with 
punctate-reticulate sculpture; ovipositor slightly 
exserted, gonostylus black. 

Female. Length: 0.85-1.1 mm (holotype 1.1 
mm). 

Head blackish, with frontovertex shining 
green, with a distinctly coppery purple sheen in 
ocellar area, interantennal prominence purplish; 
antennae yellowish, but scape and pedicel brown 
proximally on dorsal surface; pronotum blackish; 
mesoscutum and axillae similar but with weak 
blue-green sheen; scutellum shiny green or blue- 
green, basally slightly coppery; mesopleuron and 
propodeum dark purple-brown; legs, including 
coxae, totally yellow; wings hyaline, venation 
testaceous-yellow; gaster dark brown with a 
metallic green sheen, basal tergite stronger 
green-blue; ovipositor sheaths dark brown. 

Head with very fine shallow, polygonally 
reticulate sculpture on frontovertex; sculpture on 
genae longitudinally elongate; ocelli forming a 
strongly to hardly acute angle; posterior ocelli 
separated from occipital margin by about twice 
their own lengths and nearly touching eye mar- 
gins; eyes not conspicuously hairy; occipital mar- 
gin more or less rounded; frontovertex about 
one-quarter to one-sixth head width; antennae 
inserted well below ventral eye margins; anten- 
nae (Fig. 187) with pedicel plus flagellum as long 
as head width or slightly shorter; clava with 
sensory area slightly enlarged but limited to apex 
only; mandibles with one outer tooth and one 
inner broad, minutely denticulate truncation 
(Fig. 188). Relative measurements (holotype): 
head width 40, head height 37, minimum fron- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



53 



tovertex width 8, POL 3.5, OOL 0.3, eye length 
25, eye width 24, malar space 15, scape length 15, 
width 2.7; other proportions of antenna as in Fig. 
187. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum with much deeper, regular, 
punctiform-reticulate sculpture in anterior four- 
fifths or less, posterior one-fifth to one-third 
smooth and shiny; mesopleuron with shallow 
elongate reticulations; forewing with venation 
and setation at base as in Fig. 189, with only a 
very small naked basal area and basal cell with 
only a few ventral setae posteriorly; linea calva 
closed. Relative measurements (holotype): 
forewing length 90, forewing width 38; hindwing 
length 56; hindwing width 15. 

Gaster a little shorter than head plus thorax; 
ovipositor hidden or very slightly exserted; ovi- 
positor structure similar to Fig. 189; hypopygium 
similar to 25. Relative measurements (paratype): 
ovipositor length 46, gonostylus 8.5 [mid tibia 
40.5]. 

Male. Unknown. 

Variation. As noted above there is variation in 
the relative width of the frontovertex and corre- 
sponding angle formed by the ocelli and also in 
the width of the smooth part of the scutellum. 

Hosts. Unknown. 

Distribution. Malaysia, Indonesia. 

Material examined. Holotype 9> INDONE- 
SIA: Sulawesi, Utara, Dumoga-Bone NP, 
Toraut, 10.iii.1985, NH Canopy Fog #11. 
Paratypes, MALAYSIA: 1$, Sabah, Mt. Kina- 
balu NP, Poring Hot Spring, 900m, BM 
1987-349, 14.V.1987 (A. Smetana); INDONE- 
SIA: 1$, Sulawesi, Utara, Dumoga-Bone NP, 
Toraut 450m, 11. ii. 1985, BMNH Canopy Fog 
#5; 3$, Sulawesi, Utara, Dumoga-Bone NP, 
Toraut, 200m, ll.vii.1985, NH Canopy Fog #13. 
Holotype and paratypes in BMNH, paratype in 
MZB. 

Comments. The mandibular structure suggests 
that O. pilosus is close to the same group of 
species as elissa and ceres (see comments under 
elissa above). Apart from the characters given in 
the key pilosus can be separated from these 
species by having the interantennal prominence 
purplish and lacking the narrow purple line join- 
ing the ventral margins of the toruli. The other 
named species all have the interantennal promi- 
nence green and with a thin, faint purple line 
connecting the lower margins of the toruli. 
Females of O. pilosus are similar in general 



appearance to pallidipes, both species having the 
legs totally yellow, the gaster completely dark 
brown, or very nearly so, and the flagellum 
filiform, the scutellum anteriorly punctate- 
reticulate and posteriorly smooth and shiny. In 
addition to the characters given in the key pilosus 
can be separated from pallidipes by having F6 
clearly longer than broad, the forewing with only 
a very small, inconspicuous naked basal area 
with only a few ventral setae posteriorly and the 
linea calva closed. O. pallidipes has F6 more or 
less quadrate, the forewing with a conspicuous 
naked basal area with a distinct patch of ventral 
setae posteriorly and the linea calva open. 

Ooencyrtus valcanus sp.n. 

(Figs 190-192) 

Diagnosis. Female: body generally dark brown 
or blackish; head weakly metallic; mesoscutum 
faintly metallic blue; scutellum matt; all legs, 
including coxae, yellow; ovipositor sheaths dark 
brown; mandible more or less tridentate; fron- 
tovertex about one-third head width; ocelli form- 
ing a slightly acute angle; posterior ocelli 
separated from eyes by their own lengths; anten- 
nae (Fig. 190) with pedicel plus flagellum about 
1.2 times as long as head width; flagellum fili- 
form; funicle segments at least subquadrate or 
slightly longer than broad; clava with oblique 
apical truncation, as long as F3-6 together, the 
sensory part enlarged and occupying the whole of 
the ventral side of the apical segment; scutellum 
nearly flat, with regular raised reticulate sculp- 
ture and only vertical margins smooth; forewing 
(Fig. 192) without a naked basal area; linea calva 
open; postmarginal vein very short and about 
one-fourth stigmal vein; ovipositor not exserted. 

Female. Length 1 mm (holotype). 

Head blackish, frontovertex dull metallic 
green with a faint purple sheen, face and genae 
slightly more shiny, interantennal prominence 
slightly purple-blue; antennae except radicle 
yellow-testaceous; pronotum and axillae blackish 
with weak a sheen; mesoscutum weakly metallic 
blue-green; scutellum mainly matt black, poste- 
rior margin slightly metallic green, vertical mar- 
gins smooth and more strongly metallic green; 
mesopleuron and propodeum brown; legs, 
including coxae, yellow; wings hyaline, venation 
testaceous-yellow; gaster dark brown; ovipositor 
sheaths dark brown. 

Head with fairly regular, polygonally reticulate 
sculpture on frontovertex; lower parts of face 
and genae with elongate sculpture; ocelli small, 
forming a slightly acute angle; posterior ocelli 



54 



D. -W.HUANG AND J.S. NOYES 



separated from occipital margin by 1.5 times 
their own lengths and from eye margins by about 
their own lengths; eyes not conspicuously hairy; 
occipital margin not rounded; frontovertex about 
one-third head width; antennae inserted far bel- 
low the ventral eye margin; toruli separated from 
mouth margin by less than their own lengths; 
antennae (Fig. 190) with pedicel plus flagellum 
about 1.2 times as long as head width; clava 
about as long as F3-6 together, with sutures 
strongly oblique and an elongate, oblique apical 
truncation with an enlarged sensory area occupy- 
ing the whole of the truncate area; mandibles 
more or less tridentate. Relative measurements 
(holotype): head width 34, head height 30, mini- 
mum frontovertex width 12, OPL 3, POL 5, 
OOL 2, eye length 19, eye width 15, malar space 
14, scape length 18, scape width 3; other propor- 
tions of antenna as in Fig. 190. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum with deeper, regular reticu- 
late sculpture and with only vertical margins 
smooth; forewing with venation and distribution 
of setae at base as in Fig. 192, without conspicu- 
ous naked basal area and with only a few setae 
posteriorly on ventral surface of basal cell; linea 
calva open. Relative measurements (paratype): 
forewing length 98, forewing width 41; hindwing 
length 61, hindwing width 14. 

Gaster as long as thorax; ovipositor hidden, 
gonostyli about one-fifth length of ovipositor; 
hypopygium as in Fig. 191. Relative measure- 
ments (paratype): ovipositor length 40, gonosty- 
lus 8 [mid tibia 34]. 

Male. Unknown. 

Variation. Very little in material available. 

Hosts. Unknown. 

Distribution. Nepal. 

Material examined. Type material. Holotype 
$, NEPAL: Bardia, 10-24.iii.1983 (M.G. 
Allen). Paratype, 1$, same data as holotype. 
Material in BMNH. 

Comments. O. valcanus is very probably most 
closely related to mimus Prinsloo and distatus 
Prinsloo, known from the Afrotropical region. 
All three species can be characterised by the 
obliquely truncate clava and the relatively flat, 
uniformly sculptured, matt scutellum. O. valca- 
nus can be separated from these species by the 
completely yellow legs, relatively longer funicle 
segments and from distatus additionally by the 
more or less tridentate mandibles. Both mimus 
and distatus have the legs largely dark brown and 



most funicle segments distinctly transverse, 
whilst distatus has the mandibles each with two 
teeth and a clear truncation. 

Ooencyrtus urania sp.n. 

(Figs 193-195) 

Diagnosis. Female: body generally dark brown 
or blackish; head with scrobal area quite strongly 
metallic green, interantennal prominence pur- 
plish; mesoscutum with a silky appearance and 
clothed in conspicuous silvery setae; scutellum 
metallic blue-green; all legs, including coxae, 
yellow; both femora and tibia with a subapical 
brown mark adjacent to knees; gaster dark 
brown; ovipositor sheaths yellow; frontovertex 
about one-quarter head width; eyes conspicu- 
ously hairy; antennae (Fig. 193) with pedicel plus 
flagellum as long as head width; flagellum 
strongly clavate; clava slightly longer than F3-6 
together with sutures oblique and an oblique 
apical truncation, the sensory part occupying 
whole of ventral side of apical segment; mandible 
with one tooth and a broad, minutely denticulate 
truncation; scutellum slightly convex, with rela- 
tively deep sculpture anteriorly, posteriorly 
slightly shallower and only vertical margins 
smooth and shiny; forewing (Fig. 195) with only 
a very small naked basal area; linea calva closed; 
ovipositor not exserted. 

Female. Length 0.95 mm (holotype). 

Head blackish, mostly metallic green, but ocel- 
lar area coppery purple and interantennal promi- 
nence purple; antennae with radicle black, 
proximal one-third of scape yellow and distal 
two-thirds testaceous, pedicel and flagellum 
testaceous; pronotum and axillae black with a 
purple sheen; mesoscutum blackish with a weak 
green sheen and clothed in conspicuous silvery 
setae; anterior two-thirds of scutellum blue- 
green, posterior one-third similar but with a 
purplish sheen, vertical margins smooth and 
shiny, green; mesopleuron and propodeum dark 
brown; legs, including coxae, yellow; femora and 
tibiae near knees with a small brown mark; wings 
hyaline, venation testaceous-yellow; gaster dark 
brown with a metallic green sheen; ovipositor 
sheaths yellow. 

Frontovertex with fine, regular, polygonally 
reticulate sculpture which may be extremely shal- 
low between anterior ocellus and top of scrobes; 
lower face and genae with longitudinally elon- 
gate sculpture; ocelli forming an equilateral tri- 
angle; posterior ocelli separated from occipital 
margin by about 1.5 times their own lengths and 
nearly touching eye margins; eyes conspicuously 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



55 



hairy; occipital margin slightly rounded; fron- 
tovertex about one-quarter to one-fifth head 
width; head, in facial view, about 1.3 times as 
broad as high; antennae inserted just below 
ventral eye margins; toruli separated from mouth 
margin by their own lengths; antennae (Fig. 193) 
with pedicel plus flagellum about as long as head 
width; flagellum conspicuously clavate; clava 
slightly longer than F3-6 segments together, the 
sutures oblique and apex obliquely truncate with 
sensory area occupying whole of truncate part; 
mandible with one tooth and broad truncation. 
Relative measurements (holotype): head width 
44, head height 34, minimum frontovertex width 
10, OPL 4, POL 3, OOL 0.5, eye length 25, eye 
width 24, malar space 14, scape length 18, scape 
width 4; other proportions of antenna as in Fig. 
193. 

Mesoscutum with moderately deep, very fine 
imbricate-reticulate sculpture of silky appear- 
ance; scutellum moderately convex, with regular 
punctiform-reticulate sculpture in anterior two- 
thirds which is conspicuously deeper than sculp- 
ture of mesoscutum, posterior one-third with 
slightly shallower transversely reticulate sculp- 
ture, vertical part of sides and apex smooth and 
mirror shiny; forewing with venation and distri- 
bution of setae at base as in Fig. 195, without a 
naked basal area, basal cell with a few ventral 
setae posteriorly; linea calva closed. Relative 
measurements (paratype): forewing length 90, 
forewing width 41; hindwing length 62, hindwing 
width 14. 

Gaster distinctly shorter than thorax, almost 
rounded in dorsal view, last tergite apically trun- 
cate; ovipositor similar to Fig. 122 and not 
exserted; hypopygium as in 194. Relative lengths 
(paratype): ovipositor 25.5, gonostylus 5.5 [mid 
tibia 26]. 

Male. Unknown. 

Variation. There is slight variation in the rela- 
tive width of the frontovertex as outlined above. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Krakatau, Anak, 10.ix.1984 
(S. Compton). Paratypes, INDONESIA: 1$, 
Krakatau, Anak, 13. ix. 1984 (S. Compton); 1$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
Agricultural sample, BMNH Canopy Fog. #9, 
ii.1985. Holotype in MZB, paratypes in BMNH. 

Comments. Females of O. urania can be sepa- 
rated from females of other species of the genus 



with yellow legs, dark gaster and obliquely trun- 
cate clava by the conspicuously hairy eyes, 
mesoscutum with conspicuous silvery setae, 
punctate-reticulate scutellum and yellow oviposi- 
tor sheaths. 

Ooencyrtus vertumnus sp.n. 

(Figs 196-200) 

Diagnosis. Female: body generally dark 
brown; head metallic green, purplish in ocellar 
area; mesoscutum and scutellum with a metallic 
green or blue-green sheen; all legs, including 
coxae, yellow; gaster dark brown; ovipositor 
sheaths dark brown; frontovertex about one- 
quarter head width; ocelli forming a distinctly 
acute angle; posterior ocelli nearly touching 
eyes; antennae (Fig. 196) with pedicel plus flagel- 
lum as long as head width; flagellum clavate; 
funicle segments not longer than broad or hardly 
so; clava nearly as long as F3-6 together, with 
outer suture slightly oblique and apex obliquely 
truncate, the sensory area enlarged and occupy- 
ing the whole of the truncate part of the clava but 
less than one-third as long as clava; mandible 
more or less tridentate; scutellum convex, with 
regular raised punctate-reticulate sculpture but 
with veritical sides and apex smooth; forewing 
(Fig. 198) with only a small naked basal area, 
basal cell with a conspicuous narrow patch of 
ventral setae; linea calva nearly closed; postmar- 
ginal vein about as long as stigmal vein; oviposi- 
tor clearly exserted, the exserted part about as 
long as a mid tibial spur. 

Female. Length excluding ovipositor 1.05-1.22 
mm (holotype 1.17 mm). 

Ocellar area blackish with a slight purple 
sheen, scrobal area metallic green, genae with a 
slightly brassy sheen; interantennal prominence 
duller but with a metallic purple and blue line 
connecting the toruli; antennae with radicle 
black, scape yellow, pedicel and flagellum 
yellow-testaceous; pronotum, and axillae black- 
ish with a purple tinge; scutellum blue-green but 
anterior one-quarter coppery-purple, sides and 
apex shining metallic green; mesopleuron and 
propodeum dark brown; legs, including coxae, 
yellow; wings hyaline, venation testaceous- 
yellow; gaster dark brown with a slight metallic 
green sheen; ovipositor sheaths dark brown. 

Frontovertex with regular, polygonally reticu- 
late sculpture, in ocellar area moderately deep 
but shallow between anterior ocellus and top of 
scrobes; genae with longitudinally reticulate 
sculpture; ocelli relative large, forming equilat- 
eral or slightly more acute triangle; posterior 



56 



D. -W.HUANG AND J.S. NOYES 



ocelli separated from occipital margin by their 
own lengths or slightly less and nearly touching 
eye margins; eyes not conspicuously hairy; 
occipital margin rounded; frontovertex about 
one-quarter head width (Fig. 197); antennae 
inserted about far below the ventral eye margins, 
toruli about half their own lengths below eye 
margins; toruli separated from mouth margin by 
less than their own lengths; antennae (Fig. 196) 
with pedicel plus flagellum about as long as head 
width; pedicel as long as following two segments 
together; flagellum conspicuously clavate; clava 
distinctly broader than F6, and nearly as long as 
F3-6 together, the sutures parallel, but apex 
obliquely truncate, with sensory area enlarged 
and occupying the whole truncate area, about 
one-third as long as clava; mandible more or less 
tridentate in appearance. Relative measurements 
(holotype): head width 39, head height 38, mini- 
mum frontovertex width 9, OPL 3, POL 3, OOL 
0.5, eye length 26, eye width 21, malar space 16, 
scape length 17, scape width 3.5; other propor- 
tions of antenna as in Fig. 196. 

Mesoscutum with shallow, imbricate-reticulate 
sculpture; scutellum with conspicuously deeper 
punctate-reticulate sculpture, slightly shallower 
towards apex but with only the vertical sides and 
apex smooth and shiny; forewing with venation 
and distribution of setae at base as in Fig. 198, 
with only a small naked basal area, basal cell with 
a few ventral setae posteriorly; linea calva nearly 
closed. Relative measurements (paratype): 
forewing length 111, forewing width 44; hind- 
wing length 74, hindwing width 16. 

Gaster a little longer than thorax; ovipositor 
(Fig. 199) exserted; hypopygium as in Fig. 200. 
Relative lengths (paratype): ovipositor 55, gono- 
stylus 14 [mid tibia 27]. 

Male. Unknown. 

Variation. Very little variation is present in 
the material at hand, but in some specimens the 
metallic purple and blue line connecting the 
antennal toruli is inconspicuous. In the antennae 
F2-F5 may be either slightly transverse or quad- 
rate. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut 200m, BMNH Canopy Fog 
#13, ll.vii.1985. Paratypes, INDONESIA: 3$, 
Sulawesi Utara, Dumoga-Bone NP, same data as 
holotype; 1$, Sulawesi Utara, Dumoga-Bone 
NP, Toraut, 300m, 13.ii.1985, BMNH Canopy 



Fog #3; 1$, Sulawesi Utara, Dumoga-Bone NP, 
Toraut, 8.H.1985, BMNH Canopy Fog #3; 9$, 
Sulawesi Utara, Toraut, agricultural sample, 
ii.1985, BMNH Canopy Fog #9; 1$, Sulawesi 
Utara, Dumoga-Bone NP, Toraut, 10.iii.1985, 
BMNH Canopy Fog #11. Holotype in BMNH, 
paratypes in BMNH, MZB. 

Comments. Ooencyrtus vertumnus probably 
belongs to the same group of species as libitina 
and is probably closest to that species (see com- 
ments under libitina). Superficially, vertumnus is 
also similar to vesta but differs in having the 
pedicel plus flagellum as long as the head width, 
Fl-5 longer than broad or subquadrate, clava 
with sensory area about one-third as long as clava 
and scutellum with punctate-reticulate sculpture. 
In vesta the pedicel together with the flagellum is 
not more than about 0.9 times as long as head 
width, Fl-5 are strongly transverse, the sensory 
are of the clava is about half its length, and the 
scutellum has relatively shallow punctate sculp- 
ture anteriorly which is only slightly deeper than 
sculpture on meoscutum. 

Ooencyrtus vesta sp.n. 

(Figs 201, 202) 

Diagnosis. Female: body generally dark 
brown; head purplish on frontovertex, metallic 
green in scrobal area; mesoscutum with a weak 
metallic blue-green sheen; scutellum blue-green, 
basal third or so purplish; all legs, including 
coxae, yellow; gaster dark brown; ovipositor 
sheaths dark brown; frontovertex about one-fifth 
head width; antennae (Fig. 202) with pedicel plus 
flagellum distinctly shorter than head width; fla- 
gellum strongly clavate, all funicle segments 
transverse, clava with an oblique apical trunca- 
tion, as long as F2-6 together, sutures between 
segments hardly oblique, the sensory part 
enlarged and about half as long as clava; man- 
dible with three teeth; scutellum with sculpture 
in anterior half polygonally reticulate and only 
slightly deeper than sculpture on mesoscutum, 
posterior half with shallower sculpture, extreme 
sides and apex smooth and polished; forewing 
(similar to Fig. 198) with only a small naked basal 
area; linea calva closed; postmarginal vein long 
and about as long as stigmal vein; ovipositor 
(similar to Fig. 199) slightly exserted. 

Female. Length excluding ovipositor 1.1-1.2 
mm (holotype 1.13 mm). 

Frontovertex blackish with a purple sheen, 
scrobal area metallic green; interantennal promi- 
nence slightly purplish, genae a little brassy; 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



57 



antennae with radicle black, scape yellow, 
pedicel and flagellum yellow-testaceous; prono- 
tum and axillae dark purple-brown; mesoscutum 
with a weak blue-green sheen; scutellum metallic 
blue-green but coppery purple in anterior half or 
so, posterior half blue-green; mesopleuron and 
propodeum dark brown; legs, including coxae, 
yellow; wings hyaline, venation testaceous- 
yellow; gaster dark brown with a weak metallic 
blue-green sheen, especially on basal tergite; 
ovipositor sheaths dark brown. 

Frontovertex with very shallow, regularly, 
polygonally reticulate sculpture, almost smooth 
between antennal scrobes and anterior ocellus; 
genae with longitudinally elongate sculpture; 
ocelli relative large, forming an angle of about 
50°; posterior ocelli separated from occipital mar- 
gin by about 1.5 times their own lengths and 
nearly touching eye margins (Fig. 201); eyes not 
conspicuously hairy; occipital margin slightly 
rounded; frontovertex about one-fifth head 
width or a little wider; antennae inserted about 
the length of a torulus below the ventral eye 
margins; toruli separated from mouth margin by 
about half their own lengths; antennae (Fig. 202) 
with pedicel plus flagellum about 0.8-0.9 times as 
long as head width; flagellum strongly clavate 
with all funicle segments transverse; clava with 
an oblique apical truncation, about as long as 
F2-6 together, the sutures hardly oblique, the 
sensory area enlarged and about half as long as 
the clava; mandible tridentate. Relative mea- 
surements (holotype): head width 38, head 
height 37, minimum frontovertex width 8, OPL 
4, POL 3, OOL 0.3, eye length 26, eye width 23, 
malar space 15, scape length 17, scape width 3.5; 
other proportions of antenna as in Fig. 202. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum convex with regular polygo- 
nally reticulate sculpture in anterior half which is 
hardly deeper than sculpture on mesoscutum, 
posterior half with shallower, more irregular, 
elongate sculpture, the extreme apex and sides 
smooth and polished; forewing with venation and 
distribution of setae at base similar to Fig. 198, 
with only a small naked basal area, and with a 
few ventral setae posteriorly in basal cell; linea 
calva more or less closed; postmarginal about as 
long as stigmal vein. Relative measurements 
(paratype): forewing length 100, forewing width 
42; hindwing length 68, hindwing width 15. 

Gaster a little longer than thorax; ovipositor 
(similar to Fig. 199) slightly exserted; hypopy- 
gium similar to Fig. 200 but with the posterior 
margin smooth. Relative lengths (paratype): ovi- 
positor 52, gonostylus 11 [mid tibia 24]. 



Male. Unknown. 

Variation. Very little in material available. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
9, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 450m, BMNH Canopy Fog 
#5, 11. ii. 1985. Paratypes, INDONESIA: 2$, 
same data as holotype; 3$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 200m, ll.vii.1985, 
BMNH Canopy Fog #13; 29, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 8.ii.l985, BMNH 
Canopy Fog #3; 1$, Sulawesi Utara, Dumoga- 
Bone NP, Toraut, Toraut, Agricultural sample, 
ii.1985, BMNH Canopy Fog #9. Holotype in 
BMNH, paratypes in BMNH and MZBI. 

Comments. Ooencyrtus vesta is similar to mae- 
nas but differs in the sculpture of the scutellum, 
colour of the gaster and relative length of the 
ovipositor. The present species has the anterior 
part of the scutellum with sculpture that is hardly 
deeper than that on the mesoscutum, the oviposi- 
tor slightly more than twice as long as the mid 
tibia and the gaster completely dark brown. In 
maenas the scutellum has conspicuously deeper 
sculpture than the mesoscutum, the ovipositor is 
only about 1.5 times as long as the mid tibia and 
the gaster is largely yellow or orange. O. vesta is 
also superficially similar to Ooencyrtus vertum- 
nus (see comments under vertumnus, above). 

Ooencyrtus boreas sp.n. 

(Figs 203-207) 

Diagnosis. Female: head and dorsum of thorax 
bright blue-green; all legs, including coxae, yel- 
low; tegulae yellow; gaster dark brown with a 
weak metallic green sheen; ovipositor sheaths 
yellow; ocelli forming a 45° angle; occipital mar- 
gin sharp; antennae (Fig. 206) with pedicel plus 
flagellum as long as head width; flagellum fili- 
form; clava as long as F4-6 together; mandible 
with one very small tooth and a broad truncation 
which is minutely denticulate on the inner half; 
eye conspicuously hairy; frontovertex about one- 
fifth head width; scutellum slightly convex, with 
fine punctate-reticulate sculpture which is shal- 
lower near apex and sides; forewing (Fig. 207) 
with a transverse cloudy band medially and with 
a conspicuous naked basal area; linea calva open; 
postmarginal vein only slightly shorter than stig- 
mal vein; ovipositor very slightly exserted. Male: 
very similar to female but generally brighter 
metallic green; antenna with clava slightly paler 



58 



D. -W.HUANG AND J.S. NOYES 



than funicle; funicle segments quadrate or very 
slightly longer than broad and clothed in setae 
which are not longer than diameter of segments; 
mesoscutum and scutellum with very similar, 
moderately deep, fine, reticulate sculpture; 
forewing hyaline. 

Female. Length 0.95-1.10 mm (holotype 1.07 
mm). 

Head metallic blue-green with a slight violet 
sheen in ocellar area and purplish on interanten- 
nal prominence dorsally; antennae yellow, but 
radicle black; dorsum of thorax blue-green; tegu- 
lae yellow; mesopleuron and propodeum dark 
brown; legs, including coxae, yellow; wings with 
a transverse cloudy band medially from apex of 
venation, venation brown except proximal half of 
submarginal vein testaceous-yellow; gaster dark 
brown with bluish or green sheen, ovipositor 
sheaths yellow. 

Head with fairly conspicuous, almost punctate 
reticulate sculpture on frontovertex, face and 
genae; ocelli forming a 45° angle; posterior ocelli 
separated from occipital margin by about 1.5 
times their own lengths and nearly touching eye 
margins; eye conspicuously hairy; occipital mar- 
gin sharp; frontovertex about one-fifth head 
width; antennae inserted well below the lower 
eye margin; toruli separated from mouth margin 
by about their own lengths; antennae (Fig. 206) 
with pedicel plus flagellum about as long as head 
width; clava about as long as F4-6 together, with 
sensory area small and at extreme apex only; 
mandibles (Fig. 204) with one small tooth and a 
broad truncation which is minutely denticulate 
along its inner half. Relative measurements 
(holotype): head width 45, head height 40, mini- 
mum frontovertex width 9, OPL 6, POL 4, OOL 
0.3, eye length 29, eye width 26, malar space 14, 
scape length 18, scape width 4; other proportions 
of antennae as in Fig. 206. 

Mesoscutum with shallow, fine, imbricate- 
reticulate sculpture; scutellum mainly with 
deeper, almost punctate-reticulate sculpture, 
almost smooth near apex; forewing with venation 
and distribution of setae at base as in Fig. 207, 
with a distinct basal naked area, basal cell with 
only a few ventral setae posteriorly; linea calva 
open. Relative measurements (paratype): forew- 
ing length 116, forewing width 46; hindwing 
length 80, hindwing width 19. 

Gaster at most as long as thorax; ovipositor 
(Fig. 205) very slightly exserted; hypopygium as 
in Fig. 203. Relative measurements (paratype): 
ovipositor length 54, gonostylus 9 [mid tibia 55]. 

Male. Length 0.75 mm. 



Apart from genitalia, differs from female only 
in characters given in diagnosis above. 

Hosts. Reared from eggs of Stauropus lichenina 
(Lepidoptera: Notodontidae). 

Distribution. Malaysia. 

Material examined. Type material. Holotype 
$, MALAYSIA: Pahang, ex eggs of Stauropus 
lichenina Bull., 20.ii.1932 (G.H. Corbett). 
Paratypes, 9$, ld\ same data as holotype. 
Material in BMNH, IZAS. 

Comments. This species is characteristic and 
can be separated from others of the genus by the 
combination of the infuscate forewing, metallic 
blue-green head and thoracic dorsum, com- 
pletely yellow tegulae and legs and dark brown 
gaster. 

Ooencyrtus clio sp.n. 

(Figs 208-211) 

Diagnosis. Female: body generally dark 
brown; head largely metallic green or purplish; 
mesoscutum with a very weak brassy sheen; 
scutellum matt black with a very slight green 
lustre; legs, including coxae, completely yellow, 
or nearly so; gaster dark brown; ovipositor 
sheaths yellowish; ocellar area distinctly elevated 
above rest of vertex (Fig. 210); posterior ocelli 
separated from occipital margin by 2.5 times 
their own lengths; antennae (Fig. 212) with 
pedicel plus flagellum a little longer than head 
width; flagellum filiform; clava not conspicuously 
broader than F6, a little longer than F4-6 
together; mandible almost tridentate, with two 
teeth and a short truncation; scutellum almost 
entirely reticulate, with deeper sculpture than 
mesoscutum, only extreme vertical margin 
smooth and shiny; forewing (Fig. 209) hyaline 
with conspicuous naked basal area, linea calva 
open; ovipositor (similar to Fig. 266) slightly 
exserted. 

Female. Length excluding ovipositor 0.9 mm 
(holotype). 

Head with raised ocellar area dark purplish, 
rest of vertex green, slightly coppery and brassy 
anterior to ocelli; scrobal area green, lower part 
of interantennal prominence purple, genae pur- 
plish; antennae with radicle dark brown, scape 
yellow, distal half brownish dorsally; pedicel with 
proximal half brown, distal half yellow; flagellum 
yellow, and clava perhaps slightly paler than 
funicle; pronotum, mesoscutum and axillae 
blackish with weak brassy tinge; scutellum 
almost matt black but with a slight blue-green 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



59 



sheen, slightly mixed coppery near base; 
mesopleuron and propodeum dark brown; tegu- 
lae dark brown; legs, including coxae, yellow, 
but hind femur with a faint brown mark medially; 
wings hyaline, venation testaceous-yellow; gaster 
dark brown; ovipositor sheaths yellowish. 

Head with ocellar area distinctly elevated 
above rest of vertex (Fig. 210) and with fine, 
regular reticulate sculpture, postocellar area with 
rough transverse reticulate sculpture, area ante- 
rior to ocelli very smooth an shiny; ocelli forming 
an equilateral triangle; posterior ocelli separated 
from occipital margin by 2.5 times their own 
lengths and almost touching eye margins; eye 
moderately hairy; frontovertex a little less than 
one-fifth head width; occipital margin hardly 
rounded; antennae inserted just below the ven- 
tral eye margin; toruli separated from mouth 
margin by about their own lengths; antennae 
(Fig. 208) with pedicel plus flagellum slightly 
longer than head width; clava slightly longer than 
F4-6 together, with sensory area small and at 
extreme apex only; mandibles (similar to Fig. 
213) almost tridentate, with two distinct teeth 
and an emarginate inner truncation. Relative 
measurements (holotype): head width 33, head 
height 30, minimum frontovertex width 6, OPL 
7, POL 3, OOL 0, eye length 23, eye width 21, 
malar space 10, scape length 15, scape width 2.5; 
other proportions of antenna as in Fig. 208. 

Mesoscutum with moderately deep, trans- 
versely elongate, imbricate-reticulate sculpture; 
scutellum with deeper sculpture, anterior half 
medially with regular reticulate sculpture, 
towards the sides and apex conspicuously more 
elongate, but hardly shallower, only the poste- 
rior vertical margin smooth and shiny; forewing 
with venation and distribution of setae at base as 
in Fig. 209, with conspicuous naked basal area, 
basal cell with a narrow patch of ventral setae; 
linea calva open. Relative measurements 
(paratype): forewing length 79, forewing width 
36; hindwing length 54, hindwing width 13. 

Gaster longer than thorax; ovipositor similar 
to Fig. 266 and slightly exserted; hypopygium as 
in Fig. 211. Relative measurements (paratype): 
ovipositor length 38, gonostylus 8 [mid tibia 32]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut 200m, BMNH Canopy Fog 



#13, ll.vii. 1985. Paratype, 1$, same data as 
holotype. Material in BMNH. 

Comments. Ooencyrtus clio is close to circe 
both species having the ocellar area slightly to 
very distinctly raised above the rest of the vertex. 
Apart from the characters given in the key the 
two species can be separated on slight differences 
in the relative proportions of the antennal seg- 
ments, setation at the base of the forewing and 
structure of the hypopygium. In clio only Fl— 4 
are longer than broad, and F6 is slightly trans- 
verse, the forewing has a conspicuous naked 
basal area, the linea calva is open and the 
hypopygium has only a shallow posterior median 
incision, whereas in circe Fl-5 are longer than 
broad and F6 is quadrate, the forewing has no 
conspicuous basal naked area, the linea calva is 
closed and the hypopygium has a relatively deep 
posterior median incision. 

Ooencyrtus circe sp.n. 

(Figs 212-217) 

Diagnosis. Female: body generally dark 
brown; face metallic green, other parts of head 
purplish; mesoscutum slightly shiny, purple and 
blue; scutellum dark metallic green; antennae 
generally yellow with brown marks on scape and 
pedicel; wings hyaline; legs, including coxae, 
yellow; gaster dark brown; ovipositor sheaths 
dark brown; ocellar area indistinctly, but very 
slightly raised above level of frontovertex, this 
area with conspicuously rougher sculpture than 
rest of frontovertex; antennae (Fig. 212) as long 
as head width; flagellum filiform; all funicle 
segments much longer than broad; clava as long 
as F4-6 combined; mandible (Fig. 213) with a 
small outer tooth and a broad, slightly convex 
truncation; scutellum with relatively deep, 
punctate-reticulate sculpture, only the vertical 
sides and apex smooth; forewing (Fig. 214) with- 
out conspicuous naked basal area; linea calva 
closed; ovipositor very slightly exserted. 

Female. Length excluding ovipositor 0.85-1.05 
mm (holotype 1.0 mm). 

Head in ocellar area blackish dark coppery 
purple; scrobal area metallic green; interanten- 
nal prominence purplish; genae coppery; anten- 
nae with radicle black, scape yellow but distal 
half brown dorsally; pedicel proximal half 
brown, distal half yellow; flagellum yellow- 
testaceous and clava sometimes slightly paler 
than funicle; pronotum and axillae blackish with 
weak purple tinge; meoscutum relatively dull, 
blackish but with a slight purple or brassy sheen; 



60 



D. -W.HUANG AND J.S. NOYES 



scutellum dark metallic blue-green; mesopleuron 
and propodeum dark brown; tegulae dark 
brown; legs, including coxae, yellow; wings hya- 
line, venation testaceous-yellow; gaster black- 
brown; ovipositor sheaths dark brown. 

Head in ocellar area with fairly deep, polygo- 
nally reticulate sculpture which is slightly, but 
inconspicuously, raised above vertex; sculpture 
anterior to ocellar area similar but more shiny 
and almost smooth; lower parts of face and genae 
with elongate sculpture; ocelli forming an angle 
of about 50-60°; posterior ocelli separated from 
occipital margin by one and half times their own 
length and touching eye margins; eyes hairy but 
not conspicuously so; frontovertex a little less 
than one-fifth head width (Fig. 216); occipital 
margin slightly rounded; antennae inserted just 
below ventral eye margins; toruli separated from 
mouth margin by about their own lengths; anten- 
nae (Fig. 212) with pedicel plus flagellum about 
as long as head width; all funicle segments longer 
than broad, only slightly wider distally; clava 
apically rounded, as long as F4-6 together and 
with sensory part at extreme apex only; man- 
dibles (Fig. 213) with one small tooth and a 
broad, slightly concave truncation. Relative mea- 
surements (holotype): head width 38, head 
height 32, minimum frontovertex width 7, OPL 
3, POL 3, OOL 0, eye length 25, eye width 22, 
malar space 11, scape length 15, scape width 2.5; 
other proportions of antenna as in Fig. 212. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum with conspicuously deeper 
punctate-reticulate sculpture medially in anterior 
half, towards sides and apex similar, but more 
elongate, only extreme vertical sides and apex 
smooth and shiny; forewing with venation and 
distribution of setae at base as in Fig. 214, 
without conspicuous naked basal area, basal cell 
with only a few ventral setae; linea calva closed. 
Relative measurements (paratype): forewing 
length 84, forewing width 35; hindwing length 59, 
hindwing width 14. 

Gaster longer than thorax; ovipositor similar 
to Fig. 266, and at least slightly exserted; 
hypopygium as in Fig. 217. Relative measure- 
ments (paratype): ovipositor length 40, gonosty- 
lus 8 [mid tibia 34]. 

Male. Unknown. 

Variation. Very little in type material, but see 
comments below. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 



9, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #3, 
8.H.1985. Paratypes, INDONESIA: 4$, same 
data as holotype; 2$, Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #11, 
10.vii.1985; 1$, Sulawesi Utara, Dumoga-Bone 
NP, Toraut, 200m, BMNH Canopy Fog #13, 
ll.vii.1985. Holotype in BMNH, paratypes in 
BMNH and MZB. 

Non-type material. INDONESIA: 1$, BMNH 
Canopy Fog #15, 19.vii.1985; 1$, 7 Sulawesi 
Utara, Dumoga-Bone NP, Toraut, 450m, 
BMNH Canopy Fog #5, 2.H.1985. Material in 
BMNH 

Comments. The two specimens excluded from 
the paratype series differ from the type material 
in being slightly larger (1.2 mm), having the 
ocellar area markedly raised above the vertex 
(Fig. 215), the pedicel and flagellum together 
longer (about 1.15 times head width), the fron- 
tovertex a little more than one-fifth head width 
and the ovipositor about one-third longer than 
the mid tibia. These differences are generally 
very small and probably reflect a difference in 
overall body size. 

Ooencyrtus circe is closest to clio (see com- 
ments under clio above). 

Ooencyrtus clotho sp.n. 

(Figs 218-220) 

Diagnosis. Female: generally dark brown or 
blackish, but head and dorsum of thorax dis- 
tinctly metallic green or blue-green; antennae 
and all legs, including coxae, white-yellow; 
gaster dark brown; ovipositor sheaths dark 
brown; antennae with pedicel plus flagellum a 
little shorter than as head width; flagellum 
slightly clavate; funicle segments quadrate or 
transverse; head in facial view (Fig. 219) strongly 
transverse, about one-third broader than high; 
mandible almost appearing tridentate, with two 
teeth and a strongly emarginate truncation; 
mesoscutum and scutellum relatively flat; mesos- 
cutum with dense translucent, brown setae; 
scutellum with fine, regular punctiform-reticulate 
sculpture; forewing without conspicuous naked 
basal area; linea calva closed; gaster sharply 
acute apically; ovipositor slightly exserted. 

Female. Length excluding ovipositor 1.0 mm 
(holotype). 

Head in ocellar area metallic coppery purple, 
anterior to this metallic green; scrobal area 
metallic green, interantennal prominence purple, 
genae purplish with a green or blue sheen; anten- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



61 



nae with radicle pale brown, scape and pedicel 
whitish and flagellum pale yellow; pronotum and 
axillae dark purple-brown; mesoscutum metallic 
blue-green, slightly brassy posteriorly; scutellum 
metallic blue-green; mesopleuron and propo- 
deum dark brown; tegulae dark brown; legs, 
including coxae, white-yellow; wings hyaline, 
venation testaceous-yellow; gaster dark purple- 
brown with a faint metallic blue tinge; gonostyli 
dark brown. 

Head in facial view (Fig. 219) about 1.3 times 
as broad as high; frontovertex with fine, polygo- 
nally reticulate sculpture, deeper in ocellar area 
and anterior to this very shallow and almost 
smooth; lower parts of face and genae with 
elongate sculpture; ocelli forming an angle of 
about 50°; posterior ocelli separated from occipi- 
tal margin by about their own lengths and clearly 
separated from eye margins; eyes hairy but not 
conspicuously so; frontovertex a little less than 
one-quarter head width; occipital margin hardly 
rounded; antennae inserted well below ventral 
eye margin; toruli separated from mouth margin 
by about their own lengths or slightly less; anten- 
nae (similar to Fig. 251) with pedicel plus flagel- 
lum about 0.9 times as long as head width; Fl-4 
quadrate, F5-6 transverse, distal segments 
slightly larger; clava as long as F3-6 together, 
apically rounded and with sensory area small and 
restricted to apex only; mandible with two teeth 
and a strongly emarginate truncation which gives 
it an almost tridentate appearance. Relative 
measurements (holotype): head width 36, head 
height 28, minimum frontovertex width 8, OPL 
3, POL 3, OOL 1, eye length 24, eye width 20.5, 
malar space 13, scape length 15, scape width 2.5; 
other proportions of antennae similar to Fig. 251. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum flat, with distinctly deeper, 
very fine, punctate-reticulate sculpture, only 
extreme sides and apex smooth; forewing with 
venation and distribution of setae at base similar 
to Fig. 214, without conspicuous naked basal 
area basally, basal cell with only a few ventral 
setae; linea calva closed. Relative measurements 
(paratype): forewing length 76, forewing width 
35; hindwing length 55, hindwing width 15. 

Gaster longer than thorax and with apex very 
acute, pointed; ovipositor (Fig. 220) slightly 
exserted; hypopygium as in Fig. 218. Relative 
measurements (paratype): ovipositor length 50, 
gonostylus 10 [mid tibia 26]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 



Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #16, 
2.xii.l985. Paratype, INDONESIA: 1$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
200m, BMNH Canopy Fog #13, ll.vii.1985. 
Material in BMNH. 

Comments. Ooencyrtus clotho is probably close 
to clio and circe but the females can be separated 
from these and others of the genus by the combi- 
nation of very pale, whitish antennae and legs, 
strongly transverse head; flat and metallic tho- 
racic dorsum with conspicuous setae and apically 
very acute gaster. 

Ooencyrtus cybele sp.n. 

(Figs 221-225) 

Diagnosis. Female: body generally dark 
brown; head and thoracic dorsum metallic green; 
all legs, including coxae, yellow; gaster dark 
brown; antennae mostly yellowish; pedicel plus 
flagellum slightly longer than head width; flagel- 
lum filiform, all funicle segments at least slightly 
longer than broad; F2-6 much longer than broad; 
clava hardly broader than F6, as long as F4-6 
together; frontovertex about one-sixth head 
width; mandible with one tooth and broad, 
minutely denticulate truncation; scutellum with 
fine punctiform-reticulate sculpture; forewing 
(Fig. 222) with a conspicuous narrow basal naked 
area basally; linea calva open; gaster shorter than 
thorax and more or less rounded posteriorly; 
ovipositor hidden. 

Female. Length 0.8-1.15 mm (holotype 0.98 
mm). 

Head mostly metallic green, ocellar area shiny 
purplish brown; interantennal prominence pur- 
plish between toruli, ventrally delimited by a 
faint narrow bluish line; antennae testaceous- 
yellow except proximal half of pedicel brown; 
pronotum and axillae dark purple-brown; mesos- 
cutum metallic blue-green; scutellum metallic 
green; mesopleuron and propodeum blackish; 
tegulae dark brown; legs, including coxae, yel- 
low; wings hyaline, venation testaceous-yellow; 
gaster dark purple-brown with slight metallic 
green sheen on basal tergite; ovipositor sheaths 
dark brown. 

Head on frontovertex with fine, regular, 
polygonally reticulate sculpture, much shallower 
between ocelli and antennal scrobes; lower parts 
of face and genae with irregular, elongate sculp- 
ture; ocelli forming an equilateral triangle; poste- 
rior ocelli separated from occipital margin by a 



62 



D. -W.HUANG AND J.S. NOYES 



little more than their own lengths and nearly 
touching eye margins; eye not conspicuously 
hairy; frontovertex slightly more than one-sixth 
head width; occipital margin rounded; antennae 
inserted slightly below ventral eye margins (Fig. 
224); toruli separated from mouth margin by 
about their own lengths; clypeal margin medially 
slightly convex (Fig. 224); antennae (Fig. 221) 
with pedicel plus flagellum slightly longer than 
head width; all funicle segments much longer 
than broad, sometimes Fl only slightly longer 
than broad; clava hardly broader than F6, as long 
as F4-6 together, and with sensory area small 
and at extreme apex only; mandibles with one 
tooth and a broad, minutely denticulate trunca- 
tion. Relative measurements (holotype): head 
width 46, head height 40, minimum frontovertex 
width 8, OPL 4, POL 4, OOL 0.3, eye length 29, 
eye width 25.5, malar space 16, scape length 18, 
scape width 4; other proportions of antenna as in 
Fig. 221. 

Mesoscutum with regular, imbricate-reticulate 
sculpture; scutellum slightly convex with much 
deeper, more or less regular, fine, punctate- 
reticulate sculpture, only the vertical sides and 
apex smooth and polished; forewing with vena- 
tion and distribution of setae at base as in Fig. 
222, with a conspicuous narrow, naked basal 
area, basal cell with only a few ventral setae near 
base; linea calva open; postmarginal vein very 
short, less than one-third length of stigmal vein. 
Relative measurements (paratype): forewing 
length 92, forewing width 41; hindwing length 65, 
hindwing width 15. 

Gaster shorter than thorax, apex of last tergite 
truncate; ovipositor (Fig. 225) hidden; hypopy- 
gium as in Fig. 223. Relative measurements 
(paratype): ovipositor length 34, gonostylus 6 
[mid tibia 34]. 

Male. Unknown. 

Variation. Very little in material available. 

HOSTS. A parasitoid of the eggs of Riptortus sp. 
(Hemiptera: Alydidae). 

Distribution. Papua New Guinea. 

Material examined. Type material. Holotype 
?, PAPUA NEW GUINEA: Bulolo, 13.xii.1982 
(Z. Boucek). Paratypes, PAPUA NEW 
GUINEA: 1$, Bubia near Lae, 15.xii.1982 (Z. 
Boucek); 10$, Morobe Prov. Markham Valley, 
ex eggs of Riptortus sp. CIE A12822, 23.xii.1980 
(G. Young) [erroneously identified by B.R. 
Subba Rao in 1981 as Ooencyrtus erionotae Ferri- 
erel. Material in BMNH. 



Comments. The faint metallic blue line connect- 
ing the toruli, slightly produced medial part of 
the clypeal margin and minutely denticulate trun- 
cate part of the mandible suggest that O. cybele 
may be close to elissa and related species (see 
comments under elissa). It can be separated from 
these species by the characters given in the key, 
notably the completely yellow legs, unicolorous 
flagellum and uniformly reticulate scutellum. O. 
cybele is superficially similar to pallidipes but can 
be separated from this species on the dark brown 
ovipositor sheaths, sculpture of the scutellum 
and truncate apex of the last gastral tergite. In 
pallidipes the ovipositor sheaths are yellow, the 
posterior one-third or so of the scutellum is 
smooth or very nearly so and the last gastral 
tergite is apically rounded. 

Ooencyrtus ooii Noyes 

(Fig. 226) 

Ooencyrtus ooii Noyes, 1991: 1617-1622. Holo- 
type $, MALAYSIA: (BMNH, examined). 

Diagnosis. Female (length 0.8-1.15 mm): body 
generally dark brown or blackish; head metallic 
green in scrobal area; all legs, including coxae, 
yellow; mesoscutum weakly metallic green or 
blue-green, scutellum similar, anteriorly cop- 
pery; gaster dark brown; ovipositor sheaths 
honey-yellow; antennae with pedicel plus flagel- 
lum slightly longer than head width, Fl-3 quad- 
rate and clearly smaller than following three, 
F4-5 longer than broad and F6 subquadrate; 
mandible with two teeth and a truncation; scutel- 
lum convex with more or less regular punctate 
reticulate sculpture, extreme apex and sides 
smooth; forewing with a conspicuous naked basal 
area; linea calva open; postmarginal vein a little 
shorter than stigmal vein; gaster a little shorter 
than thorax; ovipositor (Fig. 226) slightly to 
distinctly exserted, about 1.5 times as long as 
mid-tibia. Male (length 0.5-0.8 mm): coloration 
similar to female but fore and hind legs often 
slightly dusky; frontovertex slightly more than 
one-third head width; antenna with Fl and F2 
the smallest, Fl quadrate, F2 slightly longer than 
broad, other funicle segments at least twice as 
long as broad and clothed in long setae more 
than twice as long as diameter of segments; 
aedeagus nearly as long as mid tibia. 

Hosts. Reared from field collected prepupae of 
Conopomorpha cramerella (Snellen) and in the 
laboratory from Acrocercops globulifera Meyrick 
(Lep.: Gracillariidae) (Ooi in Ooi et al., 1987; 
Noyes, 1991). Also recorded below from a leaf 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



63 



miner on Mangifera indica and Terminalia cat- 
appa L. 

Use in biocontrol. This species has been 
released recently in Malaysia (Sabah) for the 
control of Conopomorpha cramerella (Lepi- 
doptera: Gracillariidae) a major pest of fruit, 
especially cocoa (as Ooencyrtus sp.) (CIBC, 
1988, 1989, 1990). 

Distribution. Malaysia. 

Material examined. Type material. Holotype 
$, MALAYSIA: Sabah, Tuaran, ex pupa of 
Conopomorpha cramerella, CIBC (M) 1065, 
l.i.1988 (A.C.P. Ooi), B.M. TYPE HYM 
5,3638. Paratypes, MALAYSIA: 4$, lcf, 
Selangor, ex larva of Acrocercops globulifera, 
CIBC(M)618, 23.ix.1986 (A.C.P. Ooi); 20$, 
6cf, same data as holotype; 28$. 4cf, Sabah, 
Tuaran, lab reared, CIBC(M)1065, LL1988 
(A.C.P. Ooi); 39, 2cf, Selangor, Batu Tiga, 
gregarious parasite of larva of Acrocercops 
globulifera, CIBC(M)313, 28. ii. 1986 (A.C.P. 
Ooi); 39, Id", Selangor, Serdang, ex C. cramer- 
ella cocoon NL 311, CIBC(M)588, 22.L1986 
(A.C.P. Ooi); 119, 8cT, Selangor, Batu Tiga, lab 
host: A. globulifera, CIBC(M)1()95, 6.V.1988 
(A.C.P. Ooi); 7cf, Selangor, Batu TigA, LAB 
HOST A. globulifera, CIBC(M)1096, 7. v. 1988 
(A.C.P. Ooi); 29, Malacca, P. Gadong, ex 
Acrocercops sp., CIBC(M)1085, 13. ii. 1988 
(A.C.P. Ooi); 49, 2cf, Malacca, ex A. isonoma, 
CIBC(M)1113, 8.vii.l988 (A.C.P. Ooi). Holo- 
type in BMNH, paratypes in BMNH, TAMU, 
ZISP, USNM, PPRI, MARL 

Other material. MALAYSIA: 79, 1Q\ 
Selangor, Tg. Karang, ex Mangifera indica leaf 
miner, 28. vi. 1985, CIE A17206; 39, MARDI, ex 
Terminalia catappa L. leaf miner, CIE A 17206, 
22.11.1985. Material in BMNH. 

Comments. This species is quite similar to 
Ooencyrtus crassulus but, in addition to the char- 
acters given in the key, females of the two species 
can be separated readily by the different struc- 
ture of the antennae. In ooii F4-6 are generally 
longer than broad, whilst in crassulus F4-6 are 
quadrate. The males can be separated by the 
relative lengths of the setae on the flagellum (see 
relavant descriptions), those of crassulus being 
relatively much shorter. 

Ooencyrtus crassulus Prinsloo & Annecke 

(Fig. 227) 

Ooencyrtus crassulus Prinsloo & Annecke, 1978: 



42-44. Holotype 9, Western Samoa (ORS- 
TOM, not examined). 

Diagnosis. Female (0.6-0.8 mm): body gener- 
ally dark purple brown or blackish; head metallic 
green in scrobal area; mesoscutum with a faint 
blue sheen; scutellum matt but with apical one- 
third and sides with moderate to strong blue or 
purple lustre; all legs, including coxae, yellow; 
gaster dark brown, slightly paler basally; anten- 
nae (Fig. 227) with pedicel plus flagellum slightly 
longer than head width, Fl-3 clearly smaller than 
F4-6, which are subquadrate; mandible with one 
tooth and a broad, hardly convex truncation; 
scutellum flat, with regular, fine, relatively deep 
reticulate sculpture in anterior half or so, poste- 
rior to this shallower and more elongate, the 
vertical sides and apex smooth; forewing with 
conspicuous naked area basally, basal cell with a 
narrow patch of ventral setae; linea calva open; 
postmarginal vein a little shorter than stigmal 
vein; gaster a little shorter than thorax; oviposi- 
tor about one-quarter longer than mid-tibia, usu- 
ally hidden. Male (length 0.5-0.7 mm): 
coloration similar to female but fore and hind 
legs often slightly dusky; frontovertex about one- 
third head width; antenna with Fl and F2 the 
smallest, Fl quadrate, F2 slightly longer than 
broad, other funicle segments at least twice as 
long as broad and clothed in setae about 1.5 
times as long as diameter of segments; aedeagus 
slightly more than half as long as mid tibia. 

Hosts. Recorded by Prinsloo & Annecke ( 1978) 
as a parasitoid of the eggs of Othreis fullonia 
(Lepidoptera: Noctuidae), but also recorded 
below from eggs of Hippotion celerio L. (Lepi- 
doptera: Sphingidae). 

Distribution. Western Samoa. 

Material examined Type material. Paratype, 
19, WESTERN SAMOA: Taputimu, ex eggs of 
Othreis fullonia (Clerk), hi. 1975 (I. Swan). In 
BMNH. 

Other material. WESTERN SAMOA: 39, 
10\ Langoanca, ex eggs of Othreis fullonia, 
23.x. 1986 (D. Sands); 59, Asao, ex eggs of 
Othreis fullonia, LPL 523(A523), 20.i.l988 (W. 
Liebregts); 239, 10cf, Upolu, ex eggs of Hippo- 
tion celerio L., 10. iv. 1981 (H.J. Braune). Mate- 
rial in BMNH. 

Comments. Ooencyrtus crassulus is very close to 
sphingidarum and the two may in fact be synony- 
mous. For the present we are maintaining the 
two as distinct species because the gaster of the 
female of sphingidarum is basally yellow and the 
scutellum is strongly transverse. In crassulus the 



64 



D. -W.HUANG AND J.S. NOYES 



base of the gaster has only a slight hint of being 
paler and the scutellum is hardly broader than 
long. The antennae of the males of the two 
species are very similar. See also comments 
under ooii (above). 

Ooencyrtus lucina sp.n. (Figs 228-231) 

Diagnosis. Female: body generally yellowish; 
metanotum, propodeum and dorsal part of gaster 
dark brown; all legs, including coxae, yellow; 
gonostylus orange; frontovertex one-third head 
width; ocelli forming an equilateral triangle; 
antennae (Fig. 228) with pedicel plus flagellum as 
long as head width; funicle segments subquad- 
rate, but Fl-2 conspicuously smaller; clava 
slightly longer than F4-6 together; mandible with 
one tooth and a broad truncation; scutellum only 
slightly convex, with similar sculpture to mesos- 
cutum but posterior margin smooth; forewing 
(Fig. 229) with conspicuous naked basal area; 
linea calva open posteriorly; postmarginal vein 
about as long as stigmal vein; ovipositor not 
exserted. Male: body generally dark brown; 
scrobal area metallic green; apex of scutellum 
metallic green; scape and pedicel yellowish, Fl, 
F4 and clava dark brown, remainder of flagellum 
white; legs yellow; all funicle segments slightly 
longer than broad, longest setae about twice as 
long as diameter of segments (Fig. 231); aedea- 
gus about half as long as mid tibia. 

Female. Length 0.55-0.70 mm (holotype 0.67 
mm). 

Frontovertex orange, face yellow; antennae 
yellow but first segment of clava dark brown; 
thorax dorsally orange, sides and venter yellow; 
metanotum and propodeum dark brown; legs, 
including coxae, yellow; wings hyaline, venation 
testaceous-yellow; gaster yellow but with dor- 
sum, except basal tergite, dark brown; ovipositor 
sheaths orange. 

Head in facial view clearly transverse, about 
1.35 times as broad as high; frontovertex with 
fine, regularly raised, polygonally reticulate 
sculpture, genae with elongate sculpture; ocelli 
forming an equilateral triangle; posterior ocelli 
separated from occipital margin by their own 
lengths and distinctly separated from eye mar- 
gins; eyes not conspicuously hairy; occipital mar- 
gin quite sharp; antennae hardly inserted below 
ventral eye margins; toruli separated from mouth 
margin by less than their own lengths; antennae 
(Fig. 228) with pedicel plus flagellum about as 
long as head width; clava a little longer than F4-6 
together, with sensory area at extreme apex only; 
mandibles with one tooth and a broad trunca- 
tion. Relative measurements (holotype): head 



width 36, head height 32; minimum frontovertex 
width 12, OPL 3, POL 3, OOL 1.5, eye length 
21, eye width 16, malar space 11.5, scape length 
14, scape width 3; other proportions of antennae 
as in Fig. 228. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum mainly with similar sculp- 
ture and with only posterior vertical margin 
smooth; forewing with venation and distribution 
of setae at base as in Fig. 229, with conspicuous 
naked basal area, and basal cell with some scat- 
tered ventral setae; linea calva open; costal cell 
with only a single complete line of setae ven- 
trally. Relative measurements (paratype): forew- 
ing length 77, forewing width 34; hindwing length 
49, hindwing width 11. 

Gaster shorter than thorax; ovipositor (Fig. 
230) hidden. Relative measurements (paratype): 
ovipositor length 83, gonostylus 14 [mid tibia 71]. 

Male. Length 0.58 mm. Similar in size and leg 
coloration to female, but differs markedly in 
general body coloration, colour and structure of 
antennae, relative width of frontovertex and 
genitalia. The body is generally dark purple- 
brown with scrobal area metallic green, interan- 
tennal prominence purple and mesoscutum and 
scutellum weakly brassy, apex of scutellum 
metallic green; antennae with scape, pedicel and 
apex of clava yellow, Fl, F5 and basal half or so 
of clava brown, F2-4 and F6 white; frontovertex 
almost two-fifths head width, ocelli nearly form- 
ing a right angle, posterior ocelli about equidis- 
tant from occipital and eye margins; antennae 
(Fig. 231) with pedicel plus flagellum 1.4 times 
head width; all funicle segments slightly longer 
than broad and clothed in setae about twice as 
long as diameter of segments; aedeagus about 
half as long as mid tibia. 

Hosts. Reared from eggs of Clostera cupreata 
(Lepidoptera: Notodontidae). 

Distribution. India. 

Material examined. Type material. Holotype 
$, INDIA: Uttar Pradesh, Haldwani, ex eggs of 
Clostera cupreata, HE 22271, 3.xii.l991 (M. 
Ahmad). Paratypes, 7$, 2<3\ same data as holo- 
type. Material in BMNH. 

Comments. This is perhaps the easiest species of 
the genus to recognise. The females are remark- 
ably superficially similar to many species of 
Metaphycus, both having the body almost totally 
yellow or orange. Males, on the other hand, 
could be confused with males of some species of 
Helegonatopus , both having the flagellum simi- 
larly bicoloured. The host, mandibular structure 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



65 



and strongly expanded mesopleuron separate 
lucina from species of these other two named 
genera. Species of Metaphycus are generally 
parasitoids of scale insects, Helegonatopus are 
hyperparastioids of Dryinidae, both have triden- 
tate mandibles and the mesopleuron is not 
enlarged posteriorly and thus the metapleuron 
and propodeum are exposed laterally. 

Ooencyrtus macula sp.n. 

(Figs 232-238) 

Diagnosis Female: head metallic blue-green; 
antennae pale yellow; dorsum of thorax orange, 
laterally brown; gaster with basal half or so pale 
yellow, apex dark brown; legs, including coxae, 
pale yellow; forewing (Fig. 232) with a median 
infuscate band; ovipositor sheaths pale orange; 
mandibles with one tooth and a broad trunca- 
tion; antennae (Fig. 233) filiform with all funicle 
segments longer than broad; pedicel plus flagel- 
lum about 1.35-1.4 times head width; clava 
somewhat shorter than F4-6 together; frontover- 
tex one-sixth head width; ocelli forming a 
strongly acute angle of about 30°; forewing (Fig. 
232) with a closed, naked basal area; linea calva 
open; scutellum with fine, regular reticulate 
sculpture which is slightly deeper than that on 
mesoscutum, only posterior vertical margin 
smooth; ovipositor very slightly exserted. Male: 
similar to females but thorax generally dark 
brown, dorsum metallic green; forewing com- 
pletely hyaline; frontovertex one-third head 
width; antennae (Fig. 238) with all funicle seg- 
ments at least about 2 times as long as broad and 
clothed in setae about twice as long as diameter 
of segments. 

Female. Length 1.05-1.30 mm. (holotype 1.2 
mm). 

Head metallic blue-green on frontovertex, 
scrobal area and genae metallic green, interan- 
tennal prominence purple; radicle dark brown, 
rest of antenna pale yellow; face of pronotum 
dark brown, sides and hind margin yellowish; 
mesoscutum mainly orange but sometimes ante- 
riorly brown with a slight metallic blue sheen; 
axillae and scutellum orange; tegulae and 
prepecta pale-yellow; mesopleuron mainly 
brown but anteriorly yellowish; propodeum 
medially yellow and laterally brown; forewing 
(Fig. 232) with a transverse, median fuscous 
band; veins testaceous but proximal part of 
parastigma and marginal vein smoky; gaster 
basally pale yellow and with apical half or so dark 
purple-brown; ovipositor sheaths yellow. 

Frontovertex with very fine, regular, raised, 



polygonally reticulate sculpture; lower parts of 
face with similar sculpture but less fine, genae 
with slightly longitudinally elongate sculpture; 
ocelli forming a very acute angle of about 30°, 
posterior ocelli separated from occipital margin 
by about 1.5 times their own lengths and almost 
touching eye margins; frontovertex one-sixth 
head width; eyes with very fine but fairly con- 
spicuous setae; occipital margin hardly rounded; 
head in facial view (Fig. 234) hardly broader than 
high; antennae inserted far below the ventral eye 
margin, toruli separated from mouth margin by 
at least their own lengths; antennae (Fig. 233) 
with pedicel plus flagellum 1.35-1.4 times head 
width; clava a little shorter than F4-6 together, 
with sensory area at extreme apex only; man- 
dibles with one small tooth and a broad trunca- 
tion. Relative measurements (holotype): head 
width 42, head height 42, minimum frontovertex 
width 7, POL 2.5, OPL 4.5, OOL 0.5, eye length 
31, eye width 25, malar space 17, scape length 22, 
scape width 4.5; other proportions of antenna as 
in Fig. 233. 

Mesoscutum with shallow imbricate-reticulate, 
almost polygonally reticulate sculpture; scutel- 
lum with similar but deeper and finer regular, 
raised sculpture and with only posterior vertical 
margin smooth; mesopleuron with shallow, lon- 
gitudinally elongate reticulate sculpture; forew- 
ing with a median transverse infuscate band; 
venation and distribution of setae basally as in 
Fig. 232. Relative measurements (holotype): 
forewing length 105, forewing width 43; hind- 
wing length 69; hindwing width 15. 

Gaster conspicuously shorter than thorax; ovi- 
positor (Fig. 236) slightly exserted; hypopygium 
as in Fig. 235. Relative measurements: ovipositor 
length 72; gonostylus 15.5 [mid tibia 51]. 

Male. Length 0.78-0.92 mm. 

Similar to females except for colour of thorax, 
frontovertex width, antennae and genitalia. Tho- 
rax generally dark brown, dorsum metallic 
green, extreme sides of mesoscutum orange; 
prepecta and tegulae yellowish; forewing com- 
pletely hyaline; frontovertex one-third head 
width; antennae (Fig. 238) with pedicel plus 
flagellum 1.7 times head width, all funicle seg- 
ments at least about 2 times as long as broad and 
clothed in setae up to about twice as long as 
diameter of segments; genitalia (Fig. 237) with 
aedeagus slightly longer than half length of mid 
tibia. 

Variation. Very little in material available. 

Hosts. A gregarious parasitoid of the eggs of 



66 



D. -W.HUANG AND J.S. NOYES 



Turnaca acuta (Lepidoptera: Notodontidae) on 
coconut. 

Distribution. India. 

Material examined. Type material. Holotype 
9, INDIA: Tamil Nadu, Kamakumpeleyam, ex 
eggs of Turnaca acuta (Walker), 5.ii.l992 (S. 
Swamiappan). Paratypes, INDIA: 7$, 3d* , same 
data as holotype; 6$, 2cf, Tamil Nadu, Kalli- 
paty, ex eggs of Turnaca acuta (Walker) i. 1992 
(S. Swamiappan). Material in BMNH. 

Comments. Females of Ooencyrtus macula can 
be distinguished from other species of the genus 
because of the unique coloration, i.e. metallic 
green head contrasting with orange thoracic dor- 
sum, bicoloured gaster and infuscate forewing. 
Males should be recognised by the narrow 
orange borders to the metallic green mesoscutum 
and the bicoloured gaster. 

Ooencyrtus hera sp.n. 

(Figs 239-245) 

Diagnosis. Female: body generally dark brown 
or blackish; scrobal area metallic green; apical 
one-third and margin of scutellum shining metal- 
lic blue-green; legs, including coxae, yellow; 
gaster with at least basal tergite and venter 
orange; ovipositor sheaths dark brown, at least at 
apex; mandibles characteristic (Fig. 240), with 
one long tooth and an oblique, minutely denticu- 
late truncation; frontovertex about one-quarter 
head width (Fig. 239); flagellum (Fig. 242) fili- 
form, funicle segments all subquadrate or trans- 
verse, Fl conspicuously smaller than others; 
scutellum basally with punctate reticulate sculp- 
ture, completely smooth in posterior one-third or 
so; forewing with a conspicuous closed basal 
naked area; ovipositor very slightly exserted. 
Male: generally similar to female but frontover- 
tex about one-third head width; Fl subquadrate, 
F2-6 distinctly longer than broad (Fig. 285); 
segments clothed in setae about as long as diam- 
eter of segments. 

Female. Length, including ovipositor: 0.7-1.05 
mm (holotype 0.89 mm). 

Head purple-brown on frontovertex, scrobal 
area metallic green, interantennal prominence 
purplish; scape yellow, pedicel brownish proxi- 
mally on dorsal surface, flagellum testaceous- 
yellow; thorax blackish with a slight brassy sheen 
dorsally, apical, one-third or so scutellum shining 
blue-green; tegulae brown; legs, including coxae, 
yellow; wings hyaline, venation testaceous; 
gaster with basal tergite and venter yellow or 



testaceous, apex dorsally dark brown, sometimes 
whole gaster yellowish or testaceous; ovipositor 
sheaths dark brown, at least apically. 

Head in facial view as in Fig. 239, slightly 
broader than high; frontovertex with shallow, 
regular, raised, polygonally reticulate sculpture, 
slightly rougher in ocellar area; sculpture on 
sides of face and genae longitudinally elongate; 
ocelli forming an angle of about 50°; posterior 
ocelli separated from occipital margin by half 
their lengths and clearly separated from eyes; eye 
with conspicuous short setae; occipital margin 
sharp; antennae inserted far below the ventral 
eye margin; toruli separated from mouth margin 
by about 1.5 times their own lengths; antennae 
(Fig. 242) with pedicel plus flagellum distinctly 
shorter than head width; clava with sensory area 
at extreme apex only; mandible (Fig. 240) with 
one long tooth and an oblique, minutely denticu- 
late truncation. Relative measurements (holo- 
type); head width 43, head height 40, minimum 
frontovertex width 10, POL 3.5, OOL 0.8, eye 
length 29, eye width 23, malar space 15, scape 
length 16; other proportions of antenna as in Fig. 
242. 

Mesoscutum with shallow, almost polygonal, 
imbricate-reticulate sculpture; scutellum in ante- 
rior two thirds or so with conspicuously deeper, 
regular, punctate-reticulate sculpture, posterior 
part abruptly smooth and shiny (similar to Fig. 
106); forewing with venation and setation at base 
as in Fig. 243. Relative measurements (holo- 
type): forewing length 90, forewing width 40. 

Gaster distinctly shorter than thorax; oviposi- 
tor (Fig. 244) slightly exserted; hypopygium as in 
Fig. 241. Relative measurements (paratype): ovi- 
positor length 50, gonostylus 13.5 [mid tibia 37]. 

Male. Length 0.55-0.7 mm. Very similar to 
female but differs in coloration of the gaster, 
frontovertex width, structure of antennae and 
genitalia. Gaster totally dark brown; frontover- 
tex about one-third head width; ocelli forming an 
angle of about 80°; antennae (Fig. 245) with 
pedicel plus flagellum about 1.4 times head 
width; funicle and claval clothed with setae about 
as long as width of segment; scutellum sculptured 
area about four-fifths of scutellum; aedeagus 
about three-fifths as long as mid tibia. 

Variation. Little in material available except 
that mentioned above for coloration of female 
gaster. 



Hosts. Reared from eggs of 
glaucippe (Lepidoptera: Pieridae). 

Distribution. Thailand. 



Hebomoia 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



67 



Material examined. Type material. Holotype 
9, THAILAND: 1 1 .ii/23.ii. 1990 (Ray Harberd). 
Paratypes, THAILAND: 24$, 13d\ same data 
as holotype but various dates i-iii.1990. Material 
in BMNH, IZAS, ZISP, PPRI, USNM. 

COMMENTS. O. hera is closest to larvarum, both 
species having very similar general habitus, man- 
dibular structure with an elongate outer tooth, 
relatively long malar space, similar antennal 
structure in both sexes, ovipositor structure and 
hypopygium. The two species can be separated 
on a number of characters, but notably the 
coloration of the female gaster, relative width of 
the female frontovertex, and structure of the 
mandibles. In the female of hera the frontovertex 
is about one-quarter head width and the gaster is 
largely yellow or orange and in both sexes the 
truncate part of the mandible is oblique and 
minutely denticulate. The female of larvarum has 
the frontovertex about one-third head width, the 
gaster entirely dark brown, and in both sexes the 
mandible is tridentate. There are also differences 
in the relative proportions of the antennal seg- 
ments of both sexes (compare Figs 242, 245 and 
175, 180). In general habitus and coloration hera 
is remarkably similar to papilionis but clearly 
differs in the structure of the mandible and 
antenna. In papilionis the mandible has a broad, 
straight truncation and at least Fl and F2 are 
conspicuously smaller than the following seg- 
ments. There is also a marked difference in the 
shape of the proximal part of the second valvifer 
of the ovipositor (compare Figs 244, 322, 323). 

Ooencyrtus manii nom.nov. 

(Figs 246-254) 

Ageniaspis pyrillae Mani, 1939: 72. Holotype $, 
India (IARI, not examined). 

Ooencyrtus papilionis Ashmead; Subba Rao, 
1979: 147; 37-38; Hayat & Subba Rao, 1981: 
117; Mohyuddin et al, 1982; Asre et al., 1983: 
Dhaliwal & Bains, 1983: 294-302; Rahim & 
Hashmi, 1984: 124-126; Madan et al., 1985 
Yadhav & Chaudhary, 1984: 162-166; 1985 
949-950; Gholap & Chandele, 1985: 235-236 
Hayat, 1986;118; Chaudhary, et al. 1987 
15-20; Rajak et al., 1987: 1-9; Rahim et al., 
1991: 774-775. misidentification 

Diagnosis. Female (length about 0.65-0.95 
mm): head and thorax generally dark brown, 
gaster yellow or orange; scrobal area metallic 
green; mesoscutum weakly metallic blue; scutel- 
lum green mixed coppery; tegulae and legs, 
including coxae, yellow; gaster mainly yellow but 



margined dark brown distad of cereal plates; 
ovipositor sheaths yellow; head in facial view as 
in Fig, 247; mandible (Fig. 248) almost triden- 
tate; antennae (Figs 250, 251) with flagellum 
slightly clavate; Fl-5 a little longer than broad 
(Fig. 250), rarely transverse (Fig. 251), F6 sub- 
quadrate; clava apically rounded, as long as F4-6 
together or longer, with sensory area slightly 
enlarged but limited to apex; scutellum (Fig. 246) 
slightly convex, with regular raised, punctate- 
reticulate sculpture, only the vertical posterior 
part smooth; forewing (Fig. 249) without a naked 
basal area; linea calva open; gaster about as long 
as thorax; ovipositor (Fig. 252) hidden or only a 
little exserted, a little longer mid tibia to 1.25 
times as long as; hypopygium as in Figs 253 and 
254. Male (length about 0.8-0.9 mm): all funicle 
segments at least about twice as long as broad 
and clothed in setae a little longer than diameter 
of segments; aedeagus about half as long as mid 
tibia. 

HOSTS. A parasitoid of the eggs of Pyrilla spp., 
including the sugarcane pest, Pyrilla perpusilla 
(Homoptera: Lophopidae). 

Distribution Pakistan, India. 

Material examined. Non-type material. 
INDIA: 99, Uchani, Karnal, ex Pyrilla perpu- 
silla, CLE. A10725, ? 1978; 3$, 3cf, Punjab, 
Lyallpur, ex eggs of Pyrilla sp. 13.x. 1931; 5$, 
Punjab, Lyallpur, BM 1934-121 [identified by 
Ferriere as Ooencyrtus ?papilionis Ashmead); 
4$, Delhi, IARI area, x.1979 (Z. Boucek); 39, 
Uttar Pradesh, Gola Gokarannath, ex eggs of 
Pyrilla perpusilla, CIE A16350, x.1981 (H. 
Nigam) [identified by B.R. Subba Rao, 1979, as 
Ooencyrtus papilionis]; 2Q , Tamil Nadu, 3km E. 
Manjaler Dam, BM 1979-518, 15-18. x.1979 (J.S. 
Noyes). Material in BMNH. 

Comments. Subba Rao (1979) incorrectly syn- 
onymised Ageniaspis pyrillae Mani with Ooen- 
cyrtus papilionis Ashmead. We have examined 
the type series of papilionis and Dr M. Hayat 
(Aligarh Muslim University) has kindly supplied 
us with his notes made from examination of the 
extant type material of pyrillae in the collections 
of the Indian Agricultural Research Institute, 
Delhi. It is clear the species are not synonymous, 
differing in mandibular structure, sculpture of 
the scutellum and setation at the base of the 
forewing. In pyrillae the mandibles are triden- 
tate, the scutellum is uniformly punctate- 
reticulate with only the extreme posterior margin 
smooth and shiny and the forewing is without a 
conspicuous naked basal area. The mandibles of 



68 



D. -W.HUANG AND J.S. NOYES 



papilionis have one tooth and a broad, straight 
truncation, the scutellum has progressively 
smoother sculpture posteriorly so that the poste- 
rior two-fifths is quite shiny and the forewing has 
a conspicuous naked basal area. 

The species name pyrillae is preoccupied in 
Ooencyrtus by Ooencyrtus pyrillae Crawford, 
1916. 

Ooencyrtus manii is probably most closely 
related to midas both species having similar, 
general body coloration, mandibular structure, 
sculpture on scutellum, female clava apically 
rounded and forewing without a conspicuous 
naked basal area. Females of O. manii differs 
most significantly from those midas in coloration 
of gaster, relative lengths of flagellar segments 
and relative length of ovipositor. In manii the 
gaster is mainly yellow, but dorsally bordered 
dark brown in apical part only, the funicle seg- 
ments are subquadrate and the ovipositor is only 
slightly longer than the mid tibia and not or 
hardly exserted. O. midas has the gaster mostly 
orange-brown dorsally with only the basal tergite 
yellow, F2-6 are strongly transverse and the 
ovipositor at least twice as long as the mid tibia 
and strongly exserted, the exserted part about as 
long as the mid tibial spur. 

Ooencyrtus midas sp.n. 

(Figs 255, 259) 

Diagnosis. Female: head and thorax generally 
dark brown; head weakly metallic green in 
scrobal area and on genae; mesoscutum metallic 
blue; scutellum shining blue-green at apex and 
sides; all legs, including coxae, yellow; gaster 
orange brown dorsally, but basally and ventrally 
orange; ovipositor sheaths orange-brown; man- 
dibles tridentate; frontovertex a little less than 
one-third head width; ocelli forming an equilat- 
eral triangle; posterior ocelli clearly removed 
from eye margins; antennae (Fig. 256) with 
pedicel plus flagellum about 0.8 times head 
width; flagellar segments subquadrate or trans- 
verse; clava about as long as funicle, apex 
obliquely truncate and with sensory area 
enlarged and about half as long as clava; scutel- 
lum with regular, punctate-reticulate sculpture in 
anterior two-thirds, only posterior vertical mar- 
gin smooth and shiny; forewing (Fig. 257) with- 
out a naked basal area, gaster distinctly shorter 
than thorax, ovipositor distinctly exserted. Male: 
similar to female but antenna with all funicle 
segments subquadrate and clothed in setae a 
little longer than diameter of segments; clava 
relatively small, not wider than funicle and 



hardly longer than F5-6 together; gaster more or 
less completely brown. 

Female. Length, excluding ovipositor, about 
0.8 mm (holotype). 

Head dark purple brown in ocellar area, 
metallic green in scrobal area, interantennal 
prominence purplish; antennae with radicle 
brown, scape yellow; pedicel and flagellum testa- 
ceous; pronotum, and axillae dark purple-brown; 
meoscutum weakly metallic blue; anterior two 
thirds of scutellum weakly shining coppery 
purple, posterior one-third shining blue-green; 
mesopleuron, prepectus and propodeum dark 
purple-brown; legs, including coxae, yellow; 
wings hyaline but with a very small, indistinct 
fuscous cloud below marginal vein, venation 
testaceous; gaster dorsally orange brown with 
basal tergite and venter yellow; ovipositor 
sheaths orange-brown. 

Head with very shallow, regular, polygonally 
reticulate sculpture on frontovertex, anteriorly 
almost smooth; lower parts of face and genae 
with shallow elongate sculpture; ocelli forming 
an equilateral triangle; posterior ocelli separated 
from occipital margin by about their own lengths 
and clearly removed from eye margins; eyes 
naked; frontovertex slightly less than one-third 
head width (Fig. 255); antennae inserted well 
below ventral eye margin, toruli nearly touching 
mouth margin (Fig. 255); antennae with pedicel 
plus flagellum about 0.8 times as long as head 
width; pedicel as long as following three seg- 
ments together; flagellum distinctly clavate; clava 
collapsed in all available material but as long as 
funicle, with an oblique outer suture and almost 
certainly with an oblique apical truncation, sen- 
sory area enlarged and about half as long as 
clava; mandibles similar to Fig. 283, with three 
teeth. Relative measurements (paratype): head 
width 36, head height 30, minimum frontovertex 
width 11, POL 4, OOL 1, eye length 21, malar 
space 12, scape length 15, scape width 3.5. 

Mesoscutum with shallow almost polygonal, 
imbricate-reticulate sculpture; scutellum with 
deep regular punctate-reticulate sculpture in 
anterior two-thirds, posterior one-third becom- 
ing smoother so that apex is smooth and shiny; 
venation and distribution of setae of forewing at 
base as in Fig. 257, without a naked basal area; 
basal cell with some ventral setae posteriorly; 
linea calva nearly closed. Relative measurements 
(paratype): forewing length 90, forewing width 
37, hindwing length 60, hindwing width 12.5. 

Gaster distinctly shorter than thorax; oviposi- 
tor (Fig. 258) clearly exserted, the exserted part 
about as long as a mid tibial spur; hypopygium as 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



69 



in Fig. 259. Relative lengths (paratype): oviposi- 
tor 70, gonostylus 19.5 [mid tibia 27]. 

Male. Length about 0.8 mm. 

Similar to female but differs in coloration of 
antennae and gaster, antennal structure, fron- 
tovertex width and genitalia. Antennae yellow, 
gaster completely dark brown; funicle segments 
subquadrate and clothed in setae which are about 
as long as diameter of segments, clava hardly 
broader than F6; frontovertex very slightly wider 
than in female. 

Hosts. Reared from homopteran eggs on coco- 
nut. 

Distribution. Philippines. 

Material examined. Type material. Holotype 
$ , PHILIPPINES: Matalag, ex eggs of a uniden- 
tified homopteran en coconut, CIE A16554, 
v.1984. Paratypes, PHILIPPINES: 2$, lQ\ 
same data as holotype. Material in BMNH. 

COMMENTS. O. midas is related to libitina and 
allied species (see comments under libitina), but 
is closest to libitina. Other than differences high- 
lighted in the key to species, these two species 
differ in coloration, setation of the ventral sur- 
face of the costal cell of the forewing and struc- 
ture of the hypopygium. In midas the 
mesopleuron is dark brown, the ventral surface 
of the costal cell has only a single complete row 
of setae ventrally and the posterior margin of the 
hypopygium is smooth, whereas in libitina the 
mesopleuron is orange, the costal cell has at least 
two complete rows of setae ventrally and the 
posterior margin of the hypopygium is minutely 
serrate. O. midas may be also be close to manii 
(see comments under manii). 

Ooencyrtus mars sp.n. 

(Figs 260-264) 

Diagnosis. Female: head and thorax generally 
dark brown or blackish; scrobal area metallic 
green; mesoscutum weakly metallic green; only 
apex of scutellum shiny; all legs, including coxae, 
yellow; gaster yellowish but dorsally with apical 
tergites brown; ovipositor sheaths brown; man- 
dibles more or less tridentate; frontovertex a 
little less than one-third head width; antennae 
(Fig. 262) with all funicle segments strongly 
transverse and clava as long as funicle; sensory 
area of clava a little less than half length of clava; 
scutellum mainly with shallow sculpture, apical 
one-third smooth; ovipositor not, or hardly, 
exserted; hypopygium reaching apex of gaster or 
nearly so. 



Female. Length 0.6-0.81 mm (holotype 0.81 
mm). 

Head dark brown or blackish in ocellar area, 
scrobal area metallic green, interantennal promi- 
nence purple and genae metallic green mixed 
purple; antennae with radicle brown, scape yel- 
low; pedicel and flagellum brown; pronotum, 
mesoscutum and axillae dark brown with a weak 
metallic blue sheen; anterior two-thirds of scutel- 
lum dark purple-brown and weakly metallic, 
posterior one-third blue-green, extreme apex 
purple; mesopleuron, prepectus and propodeum 
dark brown; legs, including coxae, yellow; wings 
hyaline, venation testaceous; gaster yellow but 
with apical tergites brown; ovipositor sheaths 
brown. 

Frontovertex with relatively shallow, raised, 
polygonally reticulate sculpture; scrobal area 
with irregular, elongate sculpture; ocelli forming 
a slightly acute angle; posterior ocelli separated 
from occipital margin by at most their own 
lengths and not touching eye margins; eyes con- 
spicuously hairy; occipital margin hardly 
rounded; frontovertex a little less than one-third 
head width; head in facial view (Fig. 260) about 
1.2 times as broad as high; antennae inserted well 
below ventral eye margins; toruli separated from 
mouth margin by half their own lengths; anten- 
nae (Figs 262) with pedicel plus flagellum about 
0.8 times as long as head width; flagellum con- 
spicuously clavate; all funicle segments strongly 
transverse; clava distinctly broader than F6, and 
as long as funicle, outer suture slightly oblique 
but apex more or less rounded, sensory area 
extensive and slightly less than half as long as 
clava; mandibles similar to Fig. 268, more or less 
tridentate. Relative measurements (holotype): 
head width 32, head height 27, minimum fron- 
tovertex width 10, OPL 2, POL 4, OOL 1, eye 
length 20, eye width 16, malar space 11.5, scape 
length 12, scape width 3; other proportions of 
antenna as in Fig. 262. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum with similar, but even shal- 
lower, sculpture which gradually becomes 
weaker towards apex so that apical one-third or 
so is smooth and shiny; forewing (Fig. 261) 
without a conspicuous naked basal area; basal 
cell without ventral setae posteriorly; linea calva 
open. Relative measurements (paratype): forew- 
ing length 71, forewing width 31, hindwing length 
50, hindwing width 12. 

Gaster about as long as thorax; last tergite with 
an apical median incision; ovipositor (Fig. 264) 
not exserted or hardly so, about 1.35x as long as 
mid-tibia; hypopygium (Fig. 263) more or less 
reaching apex of gaster, and with a pair of 



70 



D. -W.HUANG AND J.S. NOYES 



submedian thickened structures anteriorly. Rela- 
tive measurements (paratype): ovipositor length 
31, gonostylus length 6 [mid tibia length 23]. 

Male. Unknown. 

Variation. Little in material available. 

Hosts. Unknown. 

Distribution. Indonesia, Papua New Guinea, 
Brunei. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 300m, BMNH Canopy Fog 
#6, 13.ii.1985. Paratypes, INDONESIA: 1$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
300m, v.1985 (J.S. Noyes); BRUNEI: 1$, Bukit 
Sulang near Lamunin, BM 1982-388, Canopy 
Fog #3, 20.viii-10.ix.1982 (N.E. Stork); 
PAPUA NEW GUINEA: 1$, Port Moresby, 
20.xii.1982 (Z. Boucek). Material in BMNH. 

Comments. Ooencyrtus mars can be separated 
from other species of the genus by the unique 
combination of strongly transverse funicle seg- 
ments with clava as long as funicle, tridentate 
mandibles, medially incised apical gastral tergite 
and hypopygium reaching apex of gaster. 

Ooencyrtus neptunus sp.n. 

(Figs 265-268) 

Diagnosis. Female: head and thorax generally 
dark brown or blackish; scrobal area metallic 
green; mesoscutum and scutellum with a slight 
metallic lustre; all legs, including coxae, yellow, 
mid femur and tibia either side of knee, each 
with a subapical dark brown ring; gaster nearly 
totally yellow but brownish along sides and at 
apex; ovipositor sheaths yellow; mandibles (Fig. 
268) tridentate; frontovertex about one-fifth 
head width; antennae with pedicel plus flagellum 
as long as head width; Fl-3 transverse and much 
smaller than F4-6; clava as long as F3-6 
together; scutellum (Fig. 267) nearly flat, with 
regular punctate-reticulate sculpture in basal 
two-thirds and only vertical margin smooth and 
shiny; forewing (Fig. 265) without a conspicuous 
naked basal area; linea calva open; ovipositor 
hidden or hardly exserted. 

Female. Length 0.65-0.85 mm (holotype 0.70 
mm). 

Frontovertex dark coppery purple to slightly 
bluish; scrobes metallic green; interantennal 
prominence purple; genae purple mixed brassy 
and green; antennae mainly yellow, dorsal side 
of scape and base of pedicel brown; pronotum 



blackish; mesoscutum and axillae blackish with a 
weak to moderately strong blue sheen; scutellum 
mainly matt black, perhaps slightly green or 
purple, vertical sides and apex shining blue- 
green; mesopleuron and propodeum dark 
brown; legs, including coxae, yellow, but mid 
legs with a pair of brown rings either side of 
knees, these sometimes also present on fore and 
hind legs; wings hyaline, venation testaceous- 
yellow; gaster mainly yellow but sides and apex 
brown; ovipositor sheaths yellow. 

Frontovertex with fairly shallow, polygonally 
reticulate sculpture, quite smooth between ante- 
rior ocellus and top of scrobes; scrobal area and 
genae with irregular, elongate sculpture; fron- 
tovertex about one-fifth head width; ocelli form- 
ing an angle of about 45-50°; posterior ocelli 
separated from occipital margin by about twice 
their own lengths and nearly touching eye mar- 
gins; eyes not conspicuously hairy; occipital mar- 
gin more or less rounded; antennae inserted well 
below ventral eye margins; toruli separated from 
mouth margin by less than their own lengths; 
antennae with pedicel plus flagellum about as 
long as head width; Fl-3 transverse and much 
smaller than F4-6; F4-5 longer than broad, F6 
subquadrate; clava slightly broader than F6, 
about as long as F3-6 together, with sensory area 
at extreme apex only; mandibles tridentate (Fig. 
268). Relative measurements (holotype): head 
width 26, head height 24, minimum frontovertex 
width 5.5, OPL 3, POL 2, OOL 0.3, eye length 
18, eye width 17, malar space 9, scape length 12, 
scape width 2.3. 

Mesoscutum with shallow, almost polygonal, 
imbricate-reticulate sculpture; scutellum (Fig. 
267) with fairly regular distinctly deeper, polygo- 
nal to elongate reticulate sculpture in basal two- 
thirds, towards apex slightly shallower and 
irregular, only vertical sides and apex smooth 
and shiny; forewing with venation and distribu- 
tion of setae at base as in Fig. 265, without 
conspicuous naked basal area, basal cell with one 
ventral row of setae posteriorly; linea calva open. 
Relative measurements (paratype): forewing 
length 69, forewing width 29; hindwing length 48 
hindwing width 10. 

Gaster slightly longer than thorax; ovipositor 
(Fig. 266) at least slightly exserted. Relative 
measurements (paratype): ovipositor length 32, 
gonostylus 6.5 [mid-tibia 25]. 

Male. Unknown. 

Variation. Very little in material available 
except that mentioned above under coloration. 

HOSTS. Unknown. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



71 



Distribution. Nepal, Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA: Sulawesi, Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy #19, 
30.ix.1985. Paratypes, NEPAL: 2$, nr Birganj, 
Lothar 550m, 12-17.ix.1967 (Can. Nepal Exp.); 
INDONESIA: 1$, Sulawesi, Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy #19, 
30. ix. 1985; 2$, Sulawesi, Utara, Dumoga-Bone 
NP, Toraut, 300m, BMNH Canopy #4, 
8.ii. 1985; 1$, Sulawesi, Utara, Dumoga-Bone 
NP, Toraut, BMNH Canopy #11, 10.iii.1985; 
6$, Sulawesi, Utara, Dumoga-Bone NP, Toraut, 
200m, BMNH Canopy #13, ll.vii.1985; 10, 
Sulawesi, Utara, Dumoga-Bone NP, Toraut, 
BMNH Canopy #15, 19.vii.1985; 2$, Sulawesi, 
Utara, Dumoga-Bone NP, Toraut, BMNH 
Canopy #16, 2.xii.l985; 1$, Sulawesi, Utara, 
Dumoga-Bone NP, Toraut, Canopy Fog, v. 1985 
(J.H. Martin). Material in BMNH, MZB, CNC. 

Comments. This species is superficially similar 
to utetheisae but differs in having tridentate man- 
dibles and brown anelli on at least the mid 
femora and tibiae. In Ooencyrtus utetheisae the 
mandibles have one tooth and a broad truncation 
and completely immaculate legs. 

Ooencyrtus libitina sp.n. 

(Figs 269-272) 

Diagnosis. Females: head and dorsum of tho- 
rax mostly metallic green or blue-green; prepec- 
tus and mesopleuron pale orange; all legs, 
including coxae, yellow; gaster pale orange but 
apical tergites dorsally brown; ovipositor sheaths 
orange to pale brown; mandibles more or less 
tridentate; frontovertex about one-quarter head 
width; flagellum (Fig. 270) clavate with funicle 
segments subquadrate or transverse; clava 
obliquely truncate, much broader than F6 and at 
least as long as F3-6 together, sensory area 
enlarged and slightly less than half length of 
clava; mandible more or less tridentate; scutel- 
lum with fairly regularly punctate-reticulate 
sculpture; ovipositor clearly exserted. 

Female. Length, excluding ovipositor, 
0.80-1.15 mm (holotype 0.96 mm). 

Head generally metallic green, ocellar area 
with purple reflections, interantennal promi- 
nence purple; antennae with radicle brown, 
scape yellow, pedicel and flagellum testaceous- 
yellow; pronotum and axillae purple-brown; 
mesoscutum metallic blue-green; scutellum 
green, only vertical margins strongly metallic; 
prepectus and mesopleuron pale orange; propo- 



deum brown; legs, including coxae, yellow; 
wings hyaline, venation testaceous-yellow; gaster 
pale-orange, dorsally with apical tergites brown; 
ovipositor sheaths orange to pale brown. 

Frontovertex with fairly shallow, regular, 
polygonally reticulate sculpture; in scrobal area 
and on genae more irregular and elongate; ocelli 
forming an equilateral triangle; posterior ocelli 
separated from occipital margin by about their 
own lengths and nearly touching eye margins; 
eyes not conspicuously hairy, clothed in numer- 
ous very fine, pale setae; occipital margin hardly 
rounded; frontovertex about one-quarter head 
width; head in facial view hardly broader than 
high; antennae inserted well bellow ventral eye 
margins; toruli separated from mouth margin by 
less than their own lengths; antennae (Fig. 270) 
with pedicel plus flagellum about as long as head 
width; funicle segments subquadrate or trans- 
verse; clava distinctly broader than F6, slightly 
longer than F3-6 together and apically obliquely 
truncate, with sensory area enlarged and only 
slightly less than half as long as clava; mandible 
more or less tridentate. Relative measurements 
(holotype): head width 33, head height 32, mini- 
mum frontovertex width 8, OPL 2.5, POL 3, 
OOL 0.5, eye length 22, eye width 18, malar 
space 12, scape length 16, scape width 3.5.; other 
proportions of antenna as in Fig. 270. 

Mesoscutum with shallow, almost polygonal, 
imbricate-reticulate sculpture; scutellum mostly 
with deeper, punctate-reticulate sculpture, more 
elongate towards sides and apex, only vertical 
sides and apex smooth and shiny; forewing with 
venation and distribution of setae at base as in 
Fig. 271, with only a small naked basal area and 
basal cell with a few ventral setae posteriorly; 
linea calva open. Relative measurements 
(paratype): forewing length 95, forewing width 
39; hindwing length 65, hindwing width 14. 

Gaster slightly longer than thorax; ovipositor 
(Fig. 272), distinctly exserted, the exserted part 
about as long as mid tibial spur; hypopygium as 
in Fig. 269, the posterior margin minutely ser- 
rate. Relative measurements (paratype): oviposi- 
tor length 61, gonostylus length 15.5 [mid tibia 
length 31]. 

Male. Unknown. 

Variation. The mesopleuron is usually pale 
orange, but in one specimen it is pale orange- 
brown. The sculpture of the flatfish dorsal part of 
the scutellum is normally more or less uniform 
but in one specimen the apex and sides are 
narrowly smooth and shiny. 

Hosts. Unknown. 



72 



D. -W.HUANG AND J.S. NOYES 



Distribution. Indonesia. 

Material examined. Type material. Holotype 
5, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 200m, BMNH Canopy Fog 
#13, ll.vii.1985. Paratypes, INDONESIA: 2$, 
same data as holotype; 2$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 200m, BMNH 
Canopy Fog #11, 10.iii.1985; 1$, Sulawesi 
Utara, Dumoga-Bone NP, Toraut, 200m, 
BMNH Canopy Fog #15, 19.vii.1985; 1$, 
Sulawesi Utara, Dumoga-Bone NP, Toraut, 
200m, FIT, ii.1985 (J.S. Noyes); 29, Sulawesi 
Utara, Dumoga-Bone NP, Toraut, 200m, v. 1985 
(J.S. Noyes); 6$, Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 200m, iv.1985 (J.S. Noyes); 
1$, Sulawesi Utara, Dumoga-Bone NP, Toraut, 
200m, 220-1 100m, iv.1985 (J.S. Noyes). Holo- 
type and paratypes in BMNH, paratypes in 
MZB, IZAS. 

Comments. O. libitina probably forms a discrete 
group of species with vertumnus, midas, luper- 
cus, lyaeus, vesta and maenas, females of these 
species having a tridentate mandible and a con- 
spicuously obliquely truncate clava. Of these 
species, libitina is probably closest to vertumnus, 
both species having the antennae situated about 
half the length of a torulus below the lowest eye 
margins, the clava conspicuously enlarged with 
sutures not strongly oblique, similar sculpture on 
the scutellum and a relatively long hypopygium 
with a minutely serrate posterior margin. O. 
libitina can be separated from vertumnus by the 
entirely orange mesopleuron, the largely orange 
gaster and the ovipositor less than twice as long 
as the mid tibia. In O. vertumnus the mesopleu- 
ron and gaster are dark brown and the ovipositor 
is at least twice as long as the mid tibia. 

O. libitina may also be close to midas (see 
comments under midas). 

Ooencyrtus lyaeus sp.n. 

(Figs 273-276) 

Diagnosis. Female: head and thorax generally 
dark brown with weak metallic reflections, 
scutellum contrasting metallic green; all legs, 
including coxae, yellow; gaster yellow-brown; 
ovipositor sheaths yellow; mandible tridentate; 
frontovertex about one-fifth head width; anten- 
nae (Fig. 275) with flagellum clavate, funicle 
segments subquadrate or transverse; clava with 
an oblique apical truncation, and as long as F3-6 
together, with sensory area enlarged but slightly 
less than half length of clava; scutellum with 
fairly uniform punctate-reticulate sculpture, 



extreme apex smooth; forewing (Fig. 276) with 
linea calva closed; ovipositor slightly exserted. 

Female. Length, excluding ovipositor, 
0.93-1.15 mm (holotype 1.05 mm). 

Frontovertex dark purple-brown, almost 
black, with a slight metallic green sheen, scrobal 
area metallic green, internatennal prominence 
purple, genae metallic green with purple reflec- 
tion; antennae with radicle brown, scape yellow, 
pedicel and flagellum testaceous-yellow; prono- 
tum and axillae dark purple-brown, mesoscutum 
similar but with a weak blue sheen; scutellum 
contrasting metallic blue-green; mesopleuron 
dark brown; propodeum brown; legs, including 
coxae, yellow; wings hyaline, venation 
testaceous-yellow; gaster brown-yellow; oviposi- 
tor sheaths pale orange. 

Ocellar area with shallow, polygonally reticu- 
late sculpture, anterior to this very shallow, 
almost smooth; scrobal area and genae with 
irregular, elongate sculpture; ocelli forming an 
angle of about 45°; posterior ocelli separated 
from occipital margin by about their own lengths 
and nearly touching eye margins; eyes not con- 
spicuously hairy; occipital margin hardly 
rounded; frontovertex about one-fifth head 
width; head in facial view only slightly broader 
than high; antennae inserted well below the 
ventral eye margin; toruli separated from mouth 
margin by about half their own lengths; antennae 
(Fig. 275) with pedicel plus flagellum about as 
long as head width; clava distinctly broader than 
F6 and as long as F3-6 together, apex obliquely 
truncate and sensory area enlarged and only 
slightly less than half length of clava; mandible 
tridentate. Relative measurements (holotype): 
head width 37, head height 35, minimum fron- 
tovertex width 7.5, OPL 3.5, POL 3, OOL 0.3, 
eye length 25, eye width 22, malar space 14, 
scape length 18, scape width 3.5; other propor- 
tions of antennae as in Fig. 275. 

Mesoscutum almost smooth but with very shal- 
low imbricate-reticulate sculpture; scutellum 
with much deeper, fairly regular, punctate- 
reticulate sculpture, only extreme sides and apex 
smooth and very shiny; forewing with venation 
and distribution of setae at base as in Fig. 276, 
without conspicuous naked basal area, basal cell 
with a patch of ventral setae posteriorly; linea 
calva closed. Relative measurements (paratype): 
forewing length 96, forewing width 39; hindwing 
length 64, hindwing width 14. 

Gaster slightly longer than thorax, almost 
rounded apically; ovipositor (Fig. 274) slightly 
exserted; hypopygium as in Fig. 273 and with 
posterior margin smooth. Relative measure- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



73 



merits (paratype): ovipositor length 49, gonosty- 
lus length 11 [mid tibia length 34]. 

Male. Unknown. 

Variation. Very little in material available. 

Hosts. Unknown. 

Distribution Indonesia. 

Material examined Holotype 9, INDONE- 
SIA: Sulawesi Utara, Dumoga-Bone NP, 
Toraut, 450m, BMNH Canopy Fog #5, 
11. ii. 1985. Paratypes, INDONESIA: 79, same 
data as holotype; 29. Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #3, 
8.ii. 1985; 39, Sulawesi Utara, Dumoga-Bone 
NP, Toraut, BMNH Canopy Fog #13, 
ll.vii. 1985. Holotype and paratypes in BMNH, 
paratypes in MZB. 

Comments. O. lyaeus belongs to the same group 
of species as libitina (see comments under lib- 
itina) and is probably closest to lupercus, maenas 
and vertumnus. It can be distinguished from 
lupercus and maenas by the presence of a distinct 
narrow patch of ventral setae posteriorly in basal 
cell; Fl-3 subquadrate and clava as long as F3-6 
and outer suture only slightly oblique. In both 
lupercus and maenas there is no conspicuous 
patch of ventral setae in basal cell posteriorly, 
Fl-3 are conspicuously transverse and the clava 
is at least as long as F2-6 together and outer 
suture conspicuously oblique. O. lyaeus differs 
from vertumnus in having the antennal toruli at 
least their own lengths below the lowest eye 
margins, eyes with short but inconspicuous setae, 
ovipositor only about 1.5 times as long as the mid 
tibia, and posterior margin of hypopygium 
smooth. In vertumnus the antennal toruli are 
only about half their own lengths below the eye 
margins, the eyes are more or less completely 
naked, the ovipositor is about twice as long as the 
mid tibia and the posterior margin of the hypopy- 
gium is minutely serrate. 

Ooencyrtus lupercus sp.n. 

(Figs 277-281) 

Diagnosis. Female: head and thorax generally 
dark brown or blackish; head and dorsum of 
thorax with a metallic green lustre; antennae 
mostly yellow with clava brown; all legs, includ- 
ing coxae, whitish or pale yellow; gaster pre- 
dominantly yellowish, apical tergites brown; 
ovipositor sheaths yellow; mandible tridentate; 
frontovertex about one-fifth head width; anten- 
nae (Fig. 277) with pedicel plus flagellum as long 



as head width; funicle segments subquadrate or 
transverse; clava as long as F2-5 together, with 
sutures oblique and a strong oblique apical trun- 
cation, sensory area occupying whole of truncate 
part; scutellum with fairly uniform punctate- 
reticulate sculpture and with only posterior mar- 
gin smooth; forewing (Fig. 268) with only a small 
basal naked area; gaster longer than thorax; 
ovipositor slightly exserted. 

Female. Length 0.9-1.0 mm (holotype 0.97 
mm). 

Head metallic green, ocellar area and interan- 
tennal prominence purplish; antennae with 
radicle black, scape yellow, pedicel proximally 
brown and distally yellowish, funicle segments 
yellowish, clava brown but with truncate area 
yellowish; pronotum and axillae black with a 
purple sheen; mesoscutum with a metallic blue- 
green sheen; reticulate area of scutellum green 
with some coppery purple reflections, posterior 
margin metallic green; prepectus and mesopleu- 
ron dark purple-brown; propodeum brown; legs, 
including coxae, pale yellow; wings hyaline, 
venation testaceous-yellow; gaster mostly yellow- 
ish, sides and apical tergites brown; ovipositor 
sheaths yellow. 

Frontovertex with fairly regular polygonally 
reticulate sculpture, a little deeper in ocellar 
area; genae with more irregular, elongate sculp- 
ture; ocelli forming an angle of about 45-50°; 
posterior ocelli separated from occipital margin 
by about 1.5 times their own lengths and nearly 
touching eye margins; eyes appearing naked; 
occipital margin slightly rounded; frontovertex 
about a little more than one-fifth head width; 
head in facial view nearly as long as broad with 
antennae inserted far below ventral eye margin, 
toruli separated from mouth margin by half their 
own lengths (Fig. 280); antennae (Fig. 277) with 
pedicel plus flagellum about as long as head 
width; clava distinctly broader than F6, about as 
long as F2-6 together, sutures oblique in profile, 
and apex distinctly obliquely truncate, with sen- 
sory part occupying whole of truncate area; man- 
dibles similar to Fig. 283 with three acute teeth. 
Relative measurements (holotype): head width 
36, head height 35, minimum frontovertex width 
7.5, OPL 4, POL 2.5, OOL 0.3, eye length 25, 
eye width 21, malar space 14, scape length 16, 
scape width 3.5; other proportions of antennae as 
in Fig. 277. 

Mesoscutum with shallow, almost polygonal, 
imbricate-reticulate sculpture; scutellum with 
conspicuously deeper, regular, punctate- 
reticulate sculpture and with only extreme apex 
and sides smooth and very shiny; forewing with 



74 



D. -W.HUANG AND J.S. NOYES 



venation and distribution of setae at base as in 
Fig. 278, and without a conspicuous naked basal 
area, basal cell without ventral setae posteriorly; 
linea calva closed. Relative measurements 
(paratype): forewing length 93, forewing width 
36; hindwing length 64, hindwing width 14. 

Gaster conspicuously longer than thorax, last 
tergite broadly truncate; ovipositor (Fig. 279) 
very slightly exserted; hypopygium as in Fig. 281. 
Relative measurements (paratype): ovipositor 
length 41, gonostylus length 8 [mid tibia length 
31]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
9, INDONESIA: Sulawesi Utara, Dumoga- 
Bone NP, Toraut, 200m, BMNH Canopy Fog 
#13, ll.vii.1985. Paratypes, INDONESIA: 4$, 
same data as holotype; 1$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 200m, 450m, BMNH 
Canopy Fog #5, 2.ii.l985; 2$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, 200m, BMNH 
Canopy Fog #15, 19.vii.1985. Holotype and 
paratypes in BMNH, paratype in MZB. 

Comments. Ooencyrtus lupercus belongs to the 
same group of species as libitina (see comments 
under libitina) and is probably closest to maenas 
and lyaeus. In addition to the characters given in 
the key, both lupercus and maenas can be sepa- 
rated from lyaeus by having F2-4 conspicuously 
transverse and clava with a strongly oblique 
outer suture, whereas in lyaeus Y2-A are sub- 
quadrate and the outer suture of the clava is only 
slightly oblique. O. lupercus can be separated 
from maenas by having the clava about 2.5 times 
as long as broad with the sensory area about 
two-thirds as long as clava, linea calva closed and 
ovipositor about 1.3 times as long as mid tibia. In 
maenas the clava is about twice as long as broad, 
the sensory part is about half as long as clava, the 
linea calva is open and the ovipositor is about 1.8 
times as long as the mid tibia. 

Ooencyrtus maenas sp.n. 

(Figs 282-285) 

Diagnosis. Head and thorax generally dark 
brown or blackish; head metallic green in scrobal 
area; mesoscutum weakly shining blue; scutellum 
metallic green mixed coppery purple; all legs, 
including coxae, pale yellow; gaster mostly yel- 
low but sides and apical tergites brown; oviposi- 
tor sheaths orange brown; mandible (Fig. 283) 



tridentate; frontovertex a little less than one- 
quarter head width; antennae (Fig. 282) with 
pedicel plus flagellum as long as head width; all 
funicle segments distinctly transverse; clava as 
long as F2-6 together, outer suture oblique, apex 
obliquely truncate and sensory area about half as 
long as clava; scutellum with uniformly punctate- 
reticulate sculpture, only extreme apex smooth; 
forewing without a distinct naked basal area; 
linea calva open; ovipositor very slightly 
exserted. 

Female. Length 0.97 mm (holotype). 

Head blackish; ocellar area purplish, anterior 
to this bluish; scrobal area metallic green, inter- 
antennal prominence purplish; genae slightly 
brassy; antennae with radicle brown, proximal 
one-third of scape brown and distal two-thirds 
yellow; pedicel mainly yellow, but marked dor- 
sally with brown in basal half; flagellum yellow; 
pronotum and axillae dark purple-brown; mesos- 
cutum blackish with metallic blue tinge; reticu- 
late area of scutellum slightly shiny coppery 
purple, sides and apex metallic green; prepectus 
and propodeum dark brown; mesopleuron dark 
purple-brown; legs, including coxae, pale yellow; 
wings hyaline, venation testaceous-yellow; gaster 
mostly yellow, its sides and apical tergites brown; 
ovipositor sheaths orange brown. 

Frontovertex area with shallow, regular, 
polygonally reticulate sculpture; lower parts of 
face and genae with irregular, elongate sculp- 
ture; ocelli forming an angle of about 45°; poste- 
rior ocelli separated from occipital margin by 
about 1.5 times their own lengths and nearly 
touching eye margins; eyes appearing naked; 
occipital margin hardly rounded; frontovertex a 
little less than one-quarter head width; head in 
front view nearly as high as broad; mandibles 
(Fig. 283) with three acute teeth; antennae 
inserted well bellow ventral eye margin; toruli 
separated from mouth margin by about half their 
own lengths; antennae (Fig. 282) with pedicel 
plus flagellum about as long as head width; 
flagellum clavate; all funicle segments distinctly 
transverse; clava distinctly broader than F6, 
about as long as F2-6 together, outer suture 
oblique, its apex obliquely truncate and sensory 
area about half as long as clava. Relative mea- 
surements (holotype): head width 32, head 
height 31, minimum frontovertex width 7.5, OPL 
3, POL 2, OOL 0.3, eye length 23, eye width 19, 
malar space 14, scape length 15, scape width 3.5; 
other proportions of antenna as in Fig. 282. 

Mesoscutum with shallow, almost polygonal, 
imbricate-reticulate sculpture; scutellum mostly 
with distinctly deeper punctate-reticulate sculp- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



75 



ture, only extreme sides and apex smooth and 
shiny; forewing without a conspicuous naked 
basal area, basal cell without ventral setae poste- 
riorly; linea calva open. Relative measurements 
(paratype): forewing length 83, forewing width 
33. 

Gaster longer than thorax, apical tergite more 
or less broadly rounded; ovipositor (Fig. 284) 
very slightly exserted; hypopygium as in Fig. 285. 
Relative measurements (paratype): ovipositor 
length 39, gonostylus length 8 [mid tibia length 
22]. 

Male. Unknown. 

Hosts. Unknown. 

Distribution. Indonesia. 

Material examined. Type material. Holotype 
$, INDONESIA; Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #3, 
8.ii.l985. Paratype, 19, same data as holotype. 
Material in BMNH. 

Comments. O. maenas is close to lupercus (see 
comments under lupercus) and vesta (see com- 
ments under vesta). 

Ooencyrtus segestes Trjapitzin 

(Figs 286-295) 

Ooencyrtus segestes Trjapitzin, 1965: 320-321, 9- 
Holotype 9- Indonesia: Komodo Island 
(ZISP) [examined]. 

Diagnosis. Macropterous and brachypterous 
forms known; head and thorax generally dark 
brown or blackish and with a metallic green 
lustre; antennae yellowish; all legs, including 
coxae, yellow; tegulae dark brown; wing vena- 
tion yellowish; gaster yellow, with apical half or 
so, dark brown; mandibles (Fig. 290) with one 
tooth and a broad, minutely denticulate trunca- 
tion; scutellum (Figs 288, 289) in basal half with 
similar sculpture to mesoscutum, apical half 
almost smooth; macropterous forms without a 
conspicuous naked basal area on forewing (Fig. 
293), linea calva open posteriorly, postmarginal 
vein hardly developed; brachypterous forms with 
forewing (Figs 291, 292) not reaching distal mar- 
gin of basal tergite of gaster. Female (length 
0.65-0.95 mm): ovipositor sheaths dark brown; 
frontovertex about one third head width (Fig. 
286); ocelli very small, separated from eyes by at 
least their own lengths; antennae (Fig. 287) fili- 
form with all funicle segments distinctly longer 
than broad; ovipositor (Fig. 294) not or hardly 
exserted. Male: similar to female but frontover- 



tex about two-fifths head width and all funicle 
segments 2-2.5 times as long as broad and 
clothed with setae about 2.5 times as long as 
diameter of segments (Fig. 295). 

The female should be recognisable from the 
above diagnosis and figures provided. The male 
is described below. 

Male. Length 0.55-0.65 mm. 

Colour as in diagnosis. 

Frontovertex with regular raised, polygonally 
reticulate sculpture; ocelli quite small, forming 
an angle of about 120°; posterior ocelli about 
equidistant from eyes and occipital margin; eyes 
almost naked; occipital margin rounded; fron- 
tovertex about three-fifths head wiuth; head in 
front view about 1.2 times as broad as high; 
antennae (Fig. 295) inserted at ventral eye mar- 
gin; toruli separated from mouth margin by 
about twice their own lengths; antennae with 
pedicel plus flagellum about 1.65 times as long as 
head width. 

Mesoscutum with shallow, imbricate-reticulate 
sculpture, slightly transversely elongate anteri- 
orly; scutellum slightly convex, with similar 
sculpture nearly to apex, but this more shallow in 
apical one-third or so; macropterous forms with 
forewing similar to Fig. 293 without a naked 
basal area, basal cell without a patch of ventral 
setae posteriorly; linea calva open; postmarginal 
vein not distinctly developed. 

Gaster slightly shorter than thorax; aedeagus 
about half as long as mid tibia. 

Hosts. Unknown, but possibly associated with 
Heteroptera in grasses since the type series was 
collected by sweeping Imperata arundinacea in 
palm savanna (Borassus) (Trjapitzin, 1965) and 
noted below from the edge of rice paddy. 

Distribution. India, Thailand, P.R. China, 
Malaysia, Indonesia. 

Material examined. Type material. Holotype 
9 (macropterous), INDONESIA: Komodo 
Island, 5.viii. 1962 (I. S. Darevsky) [one antenna 
and forewing on slide No. 1081, not examined]. 
Paratypes (brachypterous), 29, same data as 
holotype. Material in ZISP. 

Non-type material. INDIA: 119, Delhi, IARI 
area, X.1979, (Z. Boucek); 99, 5o\ Uttar 
Pradesh, Aligarh, 8-10.xi. 1979 (J. Noyes); 29, 
lCf, Karnataka, Bangalore, 19-23. ix.1979 
(J. Noyes); ld\ Karnataka, Bangalore, viii.1979 
(T. Sankaran); 19, Tamil Nadu, Siruvani Forest, 
30.ix.1979 (J. Noyes); THAILAND: 19, lcf, 
Nan Province, Nan, edge of rice paddy, 
31.viii.1985 (M.J.E. Reacher-Huber); P.R. 



76 



D. -W.HUANG AND J.S. NOYES 



CHINA: 1$, lcf, Sichuan, Huili, vi.1961 (Liao 
Dingxi); MALAYSIA: 1$, Sarawak, Long 
Lama, 13.ii.1987 (A. T. Finamore, C. Boxfield); 
INDONESIA: 8$, lcf, Sulawesi, Utara, 
Dumoga-Bone NP, Toraut, iv-v.1985, (J. 
Noyes); 1$, Sulawesi, Utara, Dumoga-Bone 
NP, Toraut, v. 1985 (A. D. Austin). Material in 
BMNH, ZISP, MZB, ZIAS, USNM, PPRI. 

Comments. O. segestes is most similar to musa 
and ferrierei (see comments under those species). 
It can be separated from similar extralimital 
species by having relatively shallow sculpture on 
the scutellum (not deeper than that on meoscu- 
tum) or the dark brown tegulae. 

Ooencyrtus ferrierei Shafee, Alam & 
Agarwal 

(Figs 296-299) 

Ooencyrtus ferrierei Shafee, Alam & Agarwal, 
1975: 97-98. Holotype $, India (ZAMU, not 
examined). 

Diagnosis. Female (length about 1-1.1 mm): 
head and thorax generally dark brown or black- 
ish with a weak to moderate metallic green sheen 
on head and dorsum of thorax; tegulae dark 
brown; all legs, including coxae, yellow; gaster 
dark brown but yellow in basal one-third or so; 
ovipositor sheaths dark brown; head in facial 
view as in Fig. 296; mandibles with a very small 
tooth and very broad, minutely denticulate inner 
truncation; antennae (Fig. 297) with pedicel plus 
flagellum as long as head width; flagellum fili- 
form, Fl-4 longer than broad, F5-6 subquad- 
rate; frontovertex about one third head width; 
scutellum at base with similar sculpture to mesos- 
cutum, gradually becoming shallower towards 
apex, extreme apex smooth and shiny; forewing 
with postmarginal vein very short, not distinctly 
developed; ovipositor (Fig. 299) hardly exserted; 
hypopygium (Fig. 298) strongly transverse. Male 
(length about 0.9 mm): generally similar to 
female but antennae with all funicle segments 
about twice as long as broad and clothed in setae 
about 1.5 times as long as diameter of segments. 

Hosts. Reared from eggs of Hemiptera on 
Solarium sp. (Shafee et al., 1975). 

Distribution. India, Indonesia. 

Material examined. Type material. 

Paratypes, 3$, INDIA: Uttar Pradesh, Nainital, 
Punthnagar, 12. vi. 1967 (A.S. Shafee). Material 
in BMNH, USNM. 

Other material. INDONESIA: 2$, Java, 
Rakata Kecil, Krakatau Cent. Expdn., 



20.ix.1984 (S.G. Compton). Material in BMNH. 

Comments. O. ferrierei is similar to macropter- 
ous specimens of segestes, both species having 
the legs completely yellow, gaster dark brown 
with basal part yellow, antennae of female more 
or less filiform, frontovertex relatively wide in 
both sexes, forewing without a conspicuous 
naked basal area, postmarginal vein of forewing 
hardly developed and similar relatively shallow 
sculpture on the scutellum. In addition to differ- 
ences in the relative lengths of the flagellum and 
funicle segments (see key to species), ferrierei 
can be separated readily from segestes by the 
relative lengths of the gonostyli (compare Figs 
294 and 299) and also in the shape of the hypopy- 
gium. In ferrierei the hypopygium has an almost 
straight posterior margin (Fig. 298) whilst in 
segestes the posterior margin of the hypopygium 
is clearly convex either side of the median inci- 
sion. 

Ooencyrtus leander sp.n. 

(Figs 300-303) 

Diagnosis. Body relatively slender. Female: 
head and thorax generally dark brown; head and 
dorsum of thorax with a metallic green or blue- 
green sheen; antennae yellow; tegulae brown; all 
legs, including coxae, yellow; gaster yellow but 
bordered dark brown; ovipositor sheaths yellow; 
frontovertex about one-fifth head width; anten- 
nae (Fig. 300) with pedicel plus flagellum slightly 
longer than head width; flagellum filiform, all 
funicle segments longer than broad; clava as long 
as F4-6 together; mandible (Fig. 303) with two, 
or perhaps three outer teeth and slightly convex, 
minutely denticulate truncation; scutellum with 
shallow sculpture in anterior half, posterior half 
smooth and shiny; forewing (Fig. 301) with a 
conspicuous narrow naked basal area basally; 
linea calva open; postmarginal vein about one- 
third length of stigmal; ovipositor slightly 
exserted. Male: generally similar to female but 
gaster mostly brown dorsally, frontovertex about 
one-third head width and antennae with all 
funicle segments at least twice as long as broad 
and clothed in setae 3-A times as long as diameter 
of segments. 

Female. Length about 0.9 mm (holotype 0.93 
mm). 

Head mainly metallic green, ocellar and post- 
ocellar area with slight coppery reflections; inter- 
antennal prominence metallic purple above 
toruli; radicle brown, rest of antennae yellow; 
pronotum and axillae dark purple-brown, weakly 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



11 



shiny; mesoscutum shining green or blue-green; 
anterior two-thirds of scutellum coppery purple, 
sides and apical one-third metallic green, slightly 
brassy; mesopleuron and propodeum dark 
purple-brown; tegulae dark brown; legs, includ- 
ing coxae, yellow; wings hyaline, venation 
testaceous-yellow; gaster almost totally yellow, 
but margined dark brown, more strongly so 
anterior to cerci; ovipositor sheaths yellow. 

Frontovertex with fine, polygonally reticulate 
sculpture; lower parts of face and genae with 
more irregular elongate sculpture; ocelli more or 
less forming an equilateral triangle; posterior 
ocelli separated from occipital margin by their 
own lengths and touching eye margins; eyes with 
short inconspicuous setae; frontovertex one- 
quarter to one-fifth head width; occipital margin 
hardly rounded; antennae inserted just below 
ventral eye margin; toruli separated from mouth 
margin by about their own lengths; clypeus very 
slightly produced medially; antennae (Fig. 300) 
filiform with pedicel plus flagellum about 1.1 
times as long as head width; clava about as long 
as F4-6 together, with sensory area at extreme 
apex only; mandibles (Fig. 303) with two or 
perhaps three outer teeth and an inner convex, 
minutely denticulate truncation. Relative mea- 
surements (holotype): head width 30, head 
height 28, minimum frontovertex width 7.5, OPL 
3, POL 3, OOL 0, eye length 21, eye width 19, 
malar space 11, scape length 12, scape width 2.5; 
other proportions of antennae as in Fig. 303. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum slightly convex, in anterior 
half with very shallow sculpture as on mesoscu- 
tum, this becoming shallower posteriorly so that 
posterior half is almost completely smooth and 
shiny; forewing with venation and distribution of 
setae at base as in Fig. 301, with a narrow oblique 
naked basal area, basal cell without ventral setae 
posteriorly; linea calva open; postmarginal vein 
about one-third stigmal vein. Relative measure- 
ments (paratype): forewing length 90, forewing 
width 34; hindwing length 64, hindwing width 13. 

Gaster as long as thorax; ovipositor (Fig. 302) 
slightly exserted; hypopygium similar to Fig. 298. 
Relative measurements (paratype): ovipositor 
length 40, gonostylus 8 [mid tibia 34]. 

Male. Length 0.9 mm. 

Generally similar to female but differs in hav- 
ing the gaster darker, relative width of frontover- 
tex, antennal structure and genitalia. Dorsum of 
gaster almost completely dark brown; antennae 
about 1.7 times as long as head width; all funicle 
segments at least twice as long as broad and 
clothed with long setae, the longest of which is 



3-4 times as long as diameter of segments; fron- 
tovertex one-third head width; ocelli forming a 
right angle, posterior ocelli separated from eye 
margins by their own lengths. 

Variation. Very little in material available 
other than that mentioned under coloration and 
frontovertex width above. 

Hosts. Unknown. 

Distribution. Papua New Guinea. 

Material examined. Type material. Holotype 
O, PAPUA NEW GUINEA: Madang Province, 
Laing, xi.1982 (P. Grootaert). Paratypes, 
PAPUA NEW GUINEA: 6£, ltf , same data as 
holotype; 19$. 5d\ Madang Province, Laing, 
20. vi. 1982 (P. Grootaert), other data same as 
holotype; 129. Madang Province, Laing, 
vii.1982 (P. Grootaert). Holotype and paratypes 
in BMNH, paratypes in IZAS, QMB, ZISP, 
IRSN, MZB. 

Comments. O. leander is very close to minor 
(Perkins) (comb.n. from Ectopiognatha) and 
major (Perkins) (comb.n. from Ectopiognatha) 
known only from Queensland, Australia. It dif- 
fers from these species in having a subcylindrical 
scape in the female and different mandibular 
structure. In both major and minor the female 
scape is distinctly broadened and fattened and 
the mandibles have four teeth. The mandibular 
structure and slightly produced clypeal margin 
also suggests that leander may be close to elissa, 
etc. (see comments under elissa), but can be 
separated from these species by lacking the faint 
metallic line connecting the ventral margins of 
the toruli and by having a largely yellow gaster. 

Ooencyrtus musa sp.n. 

(Figs 304-307) 

Diagnosis. Female: head and thorax dark 
brown or blackish with a metallic blue-green 
sheen on head and mesoscutum; antennae yel- 
low, scutellum anteriorly coppery and posteriorly 
metallic green; tegulae dark brown; all legs, 
including coxae, yellow; gaster yellow basally 
and dark brown apically; ovipositor sheaths dark 
brown; mandibles with one tooth and a broad 
straight, minutely denticulate truncation; anten- 
nae (Fig. 304) filiform, all of funicle segments 
distinctly longer than broad; frontovertex about 
one-fifth head width; anterior half of scutellum 
(Fig. 305) with shallow sculpture, posterior half 
and sides smooth and shiny; forewing without 
conspicuous naked basal area; postmarginal vein 



78 



D. -W.HUANG AND J.S. NOYES 



about one-third length of stigmal; ovipositor not 
exserted. 

Female. Length 0.78-0.85 mm (holotype 0.78 
mm). 

Head blackish with a metallic green lustre, 
ocellar area with slight coppery reflections; inter- 
antennal prominence purple; genae distinctly 
brassy; radicle black, pedicel and flagellum yel- 
low; pronotum and axillae dark purple brown, 
slightly shiny; mesoscutum metallic blue-green; 
scutellum in anterior two-thirds or so coppery 
purple, posteriorly and at sides metallic green; 
mesopleuron and propodeum dark purple- 
brown; legs, including coxae, yellow; wings hya- 
line, venation testaceous-yellow; anterior half of 
gaster yellow, posterior half dark brown, black- 
ish; ovipositor sheaths dark brown. • 

Frontovertex with regular, polygonally reticu- 
late sculpture; genae almost smooth but with 
shallow, irregular elongate sculpture; posterior 
ocelli separated from occipital margin by their 
own lengths and nearly touching eye margins; 
eyes with short inconspicuous setae; occipital 
margin hardly rounded; frontovertex about one- 
fifth head width; antennae inserted just below 
ventral eye margins; toruli separated from mouth 
margin by their own lengths; antennae (Fig. 304) 
with pedicel plus flagellum slightly longer than 
head width; flagellum filiform, all funicle seg- 
ments distinctly longer than broad; clava slightly 
longer than F4-6 together; mandible with one 
small tooth and a straight, broad, minutely den- 
ticulate truncation. Relative mea-surements 
(holotype): head width 37, head height 30, mini- 
mum frontovertex width 8, OPL 2, OPL 4, OOL 
0.3, eye length 24, eye width 20, malar space 12, 
scape length 17, scape width 3.5; other propor- 
tions of antennae as in Fig. 304. 

Mesoscutum with shallow imbricate-reticulate 
sculpture; scutellum (Fig. 305) slightly convex, 
with delicate similar, but more polygonal sculp- 
ture anteriorly which gradually becomes shal- 
lower towards apex and sides so that posterior 
one-quarter and sides are smooth; forewing with- 
out a naked basal area, basal cell without ventral 
setae; linea calva open; postmarginal vein about 
one-third length of stigmal. Relative measure- 
ments (paratype): forewing length 95, forewing 
width 40, hindwing 63, hindwing 14. 

Gaster slightly shorter than thorax; ovipositor 
(Fig. 307) hidden; hypopygium as in Fig. 306. 
Relative lengths (paratype): ovipositor 34, gono- 
stylus 6 [mid-tibia 33]. 

Male. Unknown. 

Hosts. Unknown. 



Distribution. India. 

Material examined. Type material. Holotype 
$, INDIA: Karnataka, Bannerghatta N.P., 
5.xi.l979 (Z. Boucek & J.S. Noyes). Paratypes, 
INDIA: 2$, same data as holotype. Material in 
BMNH. 

Comments. Ooencyrtus musa is generally simi- 
lar to segestes and ferrierei, all three species 
having comparable coloration, filiform antennae 
and relatively shallow sculpture on the scutellum. 
O. musa can be separated from these species by 
the relative width of the frontovertex (see key). 

Ooencyrtus telenomicida (Vassiliev) 
(Figs 308-313) 

Encyrtus telenomicida Vassiliev, 1904: 117-118. 
Syntypes, central Russia, (ZISP, not exam- 
ined). 

Schedius flavof asciatus Mercet, 1921: 315. Lecto- 
type $, Spain (IEE, examined). Synonymy 
with telenomicida by Ferriere & Voegele, 
1961: 32. 

Ooencyrtus telenomicida (Vassiliev); Romanova, 
1953: 238,240,246. 

Diagnosis. Female (length about 0.85-1.10 
mm): body compact; head and thorax dark 
brown or blackish, head metallic green usually 
with coppery, purple and bluish reflections on 
lower parts of face and especially on interanten- 
nal prominence; antennae testaceous-yellow to 
pale brown; mesoscutum metallic green with 
fairly conspicuous silvery setae; scutellum in 
basal two-thirds coppery purple, apex and sides 
metallic green; tegulae dark brown; all legs, 
including coxae, yellow; gaster blackish but yel- 
low at base; ovipositor sheaths dark brown; 
antennae (Fig. 308) with pedicel plus flagellum 
distinctly longer than head width; flagellum fili- 
form and all funicle segments much longer than 
broad; frontovertex about one-quarter to nearly 
one-third head width; head in facial view (Fig. 
312) about 1.1 times as broad as high; mandible 
with one tooth and a broad truncation; scutellum 
(Fig. 309) with longitudinally elongate reticulate 
sculpture in anterior two-thirds or so, posterior 
part smooth and shiny; forewing (Fig. 310) with 
setae in basal cell quite dense but with a small 
naked basal area; marginal vein almost quadrate 
and postmarginal vein very short; gaster slightly 
shorter than thorax; ovipositor (Fig. 311) hardly 
exserted and as long as mid-tibia; hypopygium as 
in Fig. 313. Male (length about 0.55-0.95 mm): 
generally similar to female, but gaster normally 
entirely dark brown, antennae with all funicle 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



79 



segments at least twice as long as broad and 
clothed in setae about 2.5 times as long as 
diameter of segments. 

Hosts. Recorded as a parasitoid of the eggs of 
Eurygaster integriceps (Vassiliev, 1904) and Aelia 
sp. (see Trjapitzin, 1989) (Hemiptera: Scutel- 
leridae) and Dolycoris penicillatus (Hemiptera: 
Pentatomidae) (see Myartseva, 1984). Recorded 
here as a parasitoid of eggs of Gonocerus juniperi 
on juniper (Italy) and Brachynema germarii 
(Italy). Reared in the laboratory on eggs of 
Taragama repanda (Lepidoptera: Lasiocampi- 
dae) and Amorpha populi austanti (Lepidoptera: 
Sphingidae) (see Trjapitzin, 1989). Also 
recorded here as laboratory reared from of eggs 
of Gonocerus acutangulatus (Italy). 

Distribution Throughout southern and cen- 
tral Europe, north Africa, Turkey, middle East, 
Turkmenia, Uzbekistan, Kazakhstan, central 
Russia, Pakistan. 

Material examined. Type material. Lecto- 
type 9 °f Schedius flavofasciatus Mercet, 
SPAIN: Cercedilla, L2. viii. 1916. In IEE. 

Other material. PAKISTAN: 19, Chitral 
Shenyak, 3350m. vii-viii. 1984 (W.J. Budenberg); 
1$, lcf, Hindu Kush, 3550m, viii. 1984 (W.J. 
Budenberg). Material in BMNH. 

Extralimital material. ARMENIA: 5$, 2o\ 
Sharnukh, 10.ix.1956 (V.A. Trjapitzin) [deter- 
mined by Trjapitzin as Ooencyrtus telenomicida]; 
TURKEY, BULGARIA, EGYPT, SPAIN, 
ISRAEL: 122$, 22d\ various dates and locali- 
ties. Material in BMNH. 

COMMENTS. Ooencyrtus telenomicida is a distinc- 
tive species, being characterized in the female by 
the yellow base to the gaster, mandibular struc- 
ture, filiform antennae, conspicuous silvery setae 
on the mesoscutum and sculpture of the scutel- 
lum. It is very close to, and may be synonymous 
with, two extralimital species, viz: gonoceri Vig- 
giani (from Italy) and acastus Trjapitzin (Russian 
far east and Japan). It can be separated from 
these species only on very slight differences in 
sculpture on the scutellum. 

Ooencyrtus papilionis Ashmead 

(Figs 314-328) 

Ooencyrtus papilionis Ashmead, 1905: 4-5. 
LECTOTYPE 9 (here designated), Philip- 
pines (USNM, examined). 

Ooencyrtus (Schedius) leucocerus Mercet 1922: 
150-152. Lectotype 9> Indonesia (IEE, exam- 
ined). Syn.n. 



Ooencyrtus malayensis Ferriere, 1931: 282-283. 

LECTOTYPE 9 (here designated), Malaysia 

(BMNH, examined). Syn.n. 
Ooencyrtus cochereaui Prinsloo & Annecke, 

1978: 41^12. Holotype 9, New Caledonia 

(ORSTOM, not examined). Syn.n. 

Other citations. Indian records of this species 
from Pyrilla purpusilla are erroneous and refer to 
manii (see manii). 

Diagnosis. Female (length 0.7-1.1 mm). Head 
and thorax dark brown or blackish; head metallic 
green, but frontovertex and interantennal promi- 
nence purplish; mesoscutum metallic blue or 
blue-green; scutellum blue-green to coppery 
purple in basal half or so, apex metallic blue, 
green or purple; legs, including coxae, yellow; 
gaster yellow but apical tergites yellow brown to 
dark brown; ovipositor sheaths yellow; man- 
dibles with one small tooth and a broad, very 
slightly convex truncation; antennae with proxi- 
mal two or three funicle segments transverse or 
quadrate (Figs 317-319), F4-6 normally at least a 
little longer than broad; frontovertex about one- 
sixth to one-quarter head width (Figs 314 and 
315); scutellum with shallow to moderately deep 
punctiform-reticulate sculpture anteriorly, poste- 
rior two-fifths or so smooth and shiny (Fig. 316); 
setation in basal cell of fore wing as in Figs 
320-321; gaster shorter than thorax; ovipositor 
(Figs 322 and 323) not or hardly exserted, about 
one-third longer than mid tibia; hypopygium as 
in Figs 324-326. Male (length 0.5-0.7 mm): simi- 
lar to female but generally darker and duller, 
gaster entirely brown or almost so; head in facial 
view as in Fig. 327; antennae (Fig. 328) with all 
funicle segments twice as long as broad, occa- 
sionally Fl and F2 only slightly longer than 
broad, the longest setae at least twice as long as 
diameter of segments; aedeagus about three- 
fifths as long as mid tibia. 

Hosts. Recorded below from eggs of various 
families of Lepidoptera: Papilio agamemnon (see 
also Brown, 1905), Papilio memnon, Papilio 
polytes, Papilio helenus, Papilio demoleus (see 
also Jalali & Singh, 1990), Papilio rumanzovia 
(see also Brown, 1905), Papilio aegeus (see also 
Catley, 1966), Papilio ( = Chilasa) sp., Troides 
helena (Papilionidae); Hypolimnas bolina, Kal- 
lima, Tanaecia julii, Ariadne ariadne, Phalanta 
phalantha, Junonia (= Precis) lemonias 
(Nymphalidae); Tirumala limniace, Euploea 
core, Danaus chrysippus (Danaidae); Hasora sp. 
(Hesperiidae); Heliconius charitonius (Heliconi- 
idae); Chilo terenellus (Pyralidae); Othreis fullo- 
nia (Noctuidae); Aroa cometaris (Lymantriidae); 



80 



D. -W.HUANG AND J.S. NOYES 



Cephanodes hylas (Sphingidae). The record as a 
parasitoid of the eggs of Erionota thrax (Craw- 
ford, 1911; Lepesme, 1947) may be in error for 
pallidipes. Various records of this species (as 
malayensis) reared from the eggs of Hemiptera 
(e.g. Phillips, 1941; Otanes & Sison, 1941; Goot, 
1949; Szent-Ivany & Catley, 1960; Catley, 1966; 
Sands, 1977; Young, 1982) are very probably 
misidentifications of utetheisae. 

Use in biocontrol. Ooencyrtus papilionis is 
almost certainly the species introduced, as Ooen- 
cyrtus sp., from New Caledonia into Western 
Samoa for the control of Othreis fullonia (Lepi- 
doptera: Noctuidae) (Waterhouse & Norris, 
1987). This species was supposedly introduced 
into the Solomon Islands in 1937-1938 for the 
control of Amblypelta cocophaga (Hemiptera: 
Coreidae) (Phillips, 1941) and released in Papua 
New Guinea in about 1974 for the control of 
Amblypelta theobromae (Brown) (Young, 1982). 
Coulson, et al. (1988) also recorded this species 
as having been introduced into the United States 
(Massachussets) in 1981 for the control of Anasa 
tristis (Hemiptera: Coreidae) and Nezara viridula 
(Hemiptera: Coreidae). It is likely that in these 
last three cases the species was not papilionis but 
utetheisae (see comments above). 

Distribution. India, Sri Lanka, Thailand, 
Malaysia, S. China (Guangdong, see Liao et al, 
1987: 173-174), Philippines, Indonesia, Papua 
New Guinea, New Caledonia, Vanuatu, 
Solomon Is. 

Material examined. Type material. Lecto- 
type $ of Ooencyrtus papilionis, PHILIPPINES: 
Manila (Robt. Brown) (Type No. 8125, USNM) 
[according to Ashmead the material was reared 
from eggs of Papilio sp.] (USNM). Paralectotype 
of Ooencyrtus papilionis: 1$, same data as lecto- 
type (USNM). Lectotype of Ooencyrtus leuco- 
cerus Mercet: Isle de Java (IEE). Lectotype $ of 
Ooencyrtus malayensis Ferriere, MALAYA: 
Johore, ex ova of Cephanodes hylas L., 
18.xii.1928 (G.H. Corbett), with a 'type' label, 
B.M. TYPE HYM. 5. 1,070 (BMNH). Paralecto- 
types of Ooencyrtus malayensis Ferriere, 6$, 
2d, same data as lectotype except 5.1.1929; 
10$, Id, 5.1.1929, ex host 5646, other data as 
lectotype; 2$, MALAYA: Kuala Lumpur, ex 
eggs of Papilio agamemnon L., 6. hi. 1925 (G.H. 
Corbett); 1$, ex host 5833, 14.ii.1929, other data 
as lectotype; 2$, 2d, ex. eggs of Papilio polytes 
L, l.i.1921 (W.A. Lamborn), all bearing 'co- 
type' labels (all BMNH). 

Other material. INDIA: 4$, Karnataka, 25 
km.. W. of Mudigere, B.M. 1979-518, 



28.x-3.xi. 1979 (S. Noyes); 1 $, Karnataka, 
Mudigere, B.M. 1979-518, 26.x-4.xi. 1979 (J.S. 
Noyes); 5 $, 1 d, Bangalore Karnataka, ex eggs 
of Lepidoptera, CIE 20436, ii.1988 (S. K. Jalali). 
SRI LANKA: 16$, 3d", various localities, ex 
Papilio demoleus on Citrus limon, various dates, 
xii.1989-v.1990 (HE 21373); THAILAND: 
154$, 26cf , Chedi Mae Khrua, ex Papilio mem- 
non, various dates i-vi.1990 (Ray Harberd); 
82$, 24d, Chedi Mae Khrua, ex Papilio polytes, 
various dates ii-iv.1990 (R. Harberd); 6$, Chedi 
Mae Khrua, ex Papilio helenus, various dates 
i-ii.1990 (Ray Harberd); 8$, 4d\ Chedi Mae 
Khrua, ex eggs of Papilio demoleus, various 
dates ii-iv.1990 (Ray Harberd); 30$, 6d , Chedi 
Mae Khrua, ex eggs of Papilio rumanzovia, 
various dates ii— iii. 1990 (Ray Harberd); 18$, 
5d, Chedi Mae Khrua, ex eggs of Tirumala 
limniace, various dates ii-iv.1990 (Ray Harberd); 
11$, 3d, Chedi Mae Khrua, ex eggs of Euploea 
core, various dates, ii-iv.1990 (Ray Harberd); 
8$, Chedi Mae Khrua, ex eggs of Ariadne ari- 
adne, various dates ii— ii. 1990 (Ray Harberd); 
3$, 2d", Chedi Mae Khrua, ex eggs of Phalanta 
phalantha, various dates ii — iii. 1990 (Ray Har- 
berd); 2$, Id", Chedi Mae Khrua, ex eggs of 
Precis lemonias, 10.ii/22.ii.l990 (Ray Harberd); 
2$, Id, Chedi Mae Khrua, ex eggs of Helico- 
nius charitonius (N), 4.ii/21.ii.l990 (Ray Har- 
berd); 2$, Id, Chedi Mae Khrua, ex eggs of 
Danaus chrysippus, various dates ii. 1990 (Ray 
Harberd); MALAYSIA: 4$, 2d, Penang, ex 
eggs of Papilio polytes L., 1989, CIE A20495; 
5$, Id, Penang, ex eggs of Hyp. bolina, 1989, 
CIE A20495; 3$, Penang, ex eggs of Triodes 
helena, 1989, CIE A20495; 2$, Id, Penang, ex 
eggs of Kallima sp. 15. viii. 1989, CIE A20574; 
2$, Id", Penang, ex eggs of Tanaecia julii, 
20.iii.1990, CIE A21020; 3$, Serdang UPM, 
Hasora parasite, CIE A20991; 2$, Selangor 
Batu Tiga, ex Chilasa eggs (Lep. Papilionidae), 
CIE A21064, 5.V.1988 (A.C.P. Ooi); INDONE- 
SIA, 1$, Java, Krakatau, Rakata Zwarte Hoek, 
16. ix. 1964 (S.G. Compton); 4$, Sulawesi Utara, 
Dumoga-Bone NP, Toraut, BMNH Canopy Fog 
#11, 10.iii.1985; 1$, Sulawesi Utara, Dumoga- 
Bone NP, Toraut, BMNH Canopy Fog #3, 
8.H.1985; 3$, Sulawesi Utara, Dumoga-Bone 
NP, Toraut, 450m, BMNH Canopy Fog #5, 
ll.ii.1985; 7$, Sulawesi Utara, Dumoga-Bone 
NP, Toraut, 200m, BMNH Canopy Fog #13, 
ll.vii.1985; 11$, Sulawesi, Utara, Dumoga- 
Bone NP, Toraut, iv.1985 (J.S. Noyes); PAPUA 
NEW GUINEA: 2$, lcf, Port Moresby, ex eggs 
of Papilio aegeus, ii.1961 (A. Catley); 2$, Port 
Moresby, Ramu Sugar Est., ex eggs of Chilo 
terenellus on sugarcane with Telenomus, CIE 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



81 



A20355; 1$, Bulolo, 13.xii.1982 (Z. Boucek); 
19 Areme, Musgrave R., 9.xii.l982 (Z. 
Boucek); VANUATU: 2$, Efate P.Q.S., ex 
eggs on Colocasia esculenta, 18.ix. 1987 (D. Boe); 
NEW CALEDONIA: 4 $, 3 0\ Qui Poin, ex 
eggs of Othreis fullonia, iv. 1971 (P. Cochereau) 
[det. Ooencyrtus cochereaui by D.P. Annecke); 
SOLOMON Is.: 4 $?, Honiara, ex eggs of Aroa 
cometaris, CIE A10962, 12.ii.1979 (S.J. Addi- 
son). Material in BMNH, USNM, CNC, PPRL 
ZISP, MZB, TAMU. 

Comments. We have examined material of iden- 
tified as Ooencyrtus cochereaui by Annecke from 
the same host and locality as the type series of 
this species and are confident that it must repre- 
sent a geographical race of papilionis. 

Amongst the syntypic series of Ooencyrtus 
malayensis are several specimens reared from the 
eggs of Leptocorisa acuta Thunberg (Het- 
eroptera: Coreidae) that belong to utetheisae. 

O. papilionis is very close to sphingidarum, 
shakespearei and utetheisae (see comments under 
relevant species). It is also superficially very 
similar to hera but differs strongly in mandibular 
structure. 

Ooencyrtus sphingidarum Timberlake 

Ooencyrtus sphingidarum Timberlake, 1941: 
223-225. Holotype 9' Marquesas Islands 
(BPBM, examined). 

Diagnosis. Female (length about 0.8 mm): 
body generally dark purple brown or blackish; 
head metallic green in scrobal area; mesoscutum 
with a faint blue sheen; scutellum matt but with 
apical one-third and sides with moderate to 
strong blue or purple lustre; all legs, including 
coxae, yellow; gaster dark brown, yellow basally; 
frontovertex one-quarter head width; antennae 
with ¥\-A slightly transverse and a little smaller 
than F4-6, which are quadrate; mandible with 
one tooth and a broad, hardly convex truncation; 
scutellum flat, about 1.5 times as long as broad, 
with regular fine, relatively deep reticulate sculp- 
ture, only extreme sides and apex smooth and 
shiny; linea calva open; postmarginal vein a little 
shorter than stigmal vein; gaster a little shorter 
than thorax; ovipositor hidden. Male (length 
about 0.75 mm): similar to female but frontover- 
tex nearly half head width; antenna with Fl 
subquadrate, F2-6 clearly longer than broad and 
clothed in setae about 1.5 times as long as 
diameter of segments. 

Hosts. According to Timberlake (1941) the type 
series was reared from sphingid eggs (probably 



Chromis erotus eras) on Morinda citrifolia. 

Distribution. Marquesas Islands. 

Material examined. Type material. Holotype 
9, MARQUESAS IS: Nukuhiva, ex sphingid 
eggs, Nov. 25. 29 (Mumford and Adamson). 
Paratype, Id", same data as holotype. Material 
in BPBM. 

Comments. O. sphingidarum is very close to, 
and may be synonymous with, both papilionis 
and crassulus. It differs from papilionis in having 
the scutellum strongly transverse and longest 
setae on the male antennae only about 1.5 times 
as long as diameter of segments. In papilionis the 
scutellum is about one-third broader than long 
and the longest setae on the male antenna are at 
least twice as long as diameter of segments. For 
differences from crassulus see comments under 
that species. 

Ooencyrtus shakespearei (Girault) 

Coccidoxenus shakespearei Girault, 1923: 48. 

LECTOTYPE 9 (here designated), Australia 

(QMB, examined). 
Ooencyrtus shakespearei (Girault) Noyes & 

Hayat, 1984: 309. 

Diagnosis. Female (length 0.7-0.97 mm): head 
and thorax generally dark brown; head metallic 
green, interantennal prominence purple, genae 
sometimes coppery; antennae yellow; mesoscu- 
tum metallic blue; tegulae yellow in basal half; 
scutellum relatively dull, coppery purple, its mar- 
gins shining green or blue-green; all legs, includ- 
ing coxae, yellow; basal half or so of gaster 
yellow, apex dark brown; ovipositor sheaths yel- 
low; mandibles with one tooth and a broad, 
slightly convex truncation; frontovertex less than 
one-fifth head width; antennae filiform with 
pedicel plus flagellum about 1.1-1.2 times as long 
as head width; all funicle segments much longer 
than broad; clava hardly broader than F6, as long 
as F4-6 together; scutellum only slightly convex, 
with regular fine, punctate-reticulate sculpture 
which is distinctly deeper than imbricate- 
reticulate sculpture on mesoscutum, only 
extreme sides and apex smooth and shiny; forew- 
ing with a conspicuous closed, naked basal area; 
linea calva open; postmarginal vein about half 
stigmal vein; ovipositor very slightly exserted. 

Hosts. Recorded below as a hyperparasite of 
Agonoxena pyrogramma (Lepidoptera: Agonox- 
enidae). 

Distribution. Thailand, Indonesia, Papua New 
Guinea, Australia. 



82 



D. -W.HUANG AND J.S. NOYES 



Material examined. Type material. Lecto- 
type $, on slide under coverslip nearest label 
'Coccidoxenus shakespearei Type', with syntype 
of Mirsyrpophagus columbi and fragments of 
Anagyropsis spongitus Girault; according to 
Girault, 1923 the specimen was one of a pair 
collected at Nelson, Queensland, Australia 
(QMB). 

Non type material. THAILAND: 1$, Huai 
Kha Khaeng, ii.1986 (M.G. Allen); INDONE- 
SIA: 19, Sulawesi: Utara, Dumoga-Bone NP, 
Toraut, iv.1985 (J.S. Noyes); PAPUA NEW 
GUINEA: 3$, Lae, Hyper, of A pyrogramma, 
1957 (R.W. Paine). Material in BMNH. 

Extralimital material. AUSTRALIA: 1$, 
Queensland, Tibrogargan, 5.vi.l980 (J.S. Noyes) 
[compared with lectotype]. In BMNH. 

Comments. O. shakespearei is very similar in 
general habitus and coloration to papilionis and 
utetheisae. It can be separated from both species 
on the yellow tegulae (although some specimens 
of papilionis may have tegulae which are orange 
brown basally) , from papilionis by the relatively 
longer funicle segments and from utetheisae by 
the paler yellow or orange ovipositor sheaths. 
Otherwise the three species can be separated 
reliably on the characters given in the key. 

Ooencyrtus utetheisae (Risbec) 
(Figs 329-337) 

Aenasioidea utetheisae Risbec 1951: 141-143. 

Syntypes $cf, Ivory Coast (MNHN, not 

examined). 
Ooencyrtus patriciae Subba Rao, 1981: 39-41. 

Holotype $, Nigeria (BMNH, examined). 

Synonymy with utetheisae by Prinsloo, 1987: 

9-10. 
Ooencyrtus utetheisae (Risbec); Prinsloo, 1987: 

9-10. 

Diagnosis. Female (length 0.75-1.22 mm): 
head and thorax generally dark brown or black- 
ish; head weakly metallic green or blue, purplish 
on interantennal prominence; mesoscutum 
metallic blue; scutellum dark coppery purple on 
reticulate part, extreme apex and sides greenish; 
all legs, including coxae, yellow; gaster yellow or 
orange bordered dorsally dark brown or black- 
ish; outer plates of ovipositor and ovipositor 
sheaths dark brown; mandibles with a small 
outer tooth and a broad minutely denticulate 
truncation (Fig. 333) or without an outer tooth 
(Fig. 334); frontovertex about one-quarter head 
width or narrower; antennae (Figs 329 and 331) 
more or less filiform with pedicel plus flagellum 



slightly longer than head width; all funicle seg- 
ments longer than broad; clava hardly broader 
than F6, as long as F4-6 together; scutellum only 
slightly convex, with fine regular punctate- 
reticulate sculpture extreme sides and apex 
smooth; forewing (Figs 330 and 332) without a 
conspicuous naked basal area, and basal cell 
without distinct ventral setae posteriorly; linea 
calva open; postmarginal vein about half stigmal 
vein; ovipositor (Fig. 335) hidden, or hardly 
exserted, about as long as mid-tibia; hypopygium 
as in Fig. 336. Male (length 0.55-0.95 mm): 
generally similar to female, but frequently more 
strongly metallic green on head and thoracic 
dorsum; gaster totally brown, occasionally yel- 
lowish towards base; frontovertex two-fifths head 
width; antennae (Fig. 337) with pedicel and 
flagellum together about 1.8 times head width, 
all funicle segments at least twice as long as 
broad and longest setae at least twice as long as 
diameter of segments; aedeagus about half as 
long as mid tibia. 

Hosts. The type series of utetheisae was 
recorded, perhaps erroneously, as having been 
reared from the eggs of Utetheisa pulchella (Lepi- 
doptera: Arctiidae), but otherwise the species 
seems to be restricted to heteropterous eggs. In 
Africa, Prinsloo (1987) records the species from 
the eggs of Anoplocnemis curvipes, Pseudother- 
aptus wayi, Clavigralla elongata, C. tomentosicol- 
lis (Coreidae), Mirperus jaculus and Riptortus 
dentipes (Alydidae). Recorded below from the 
eggs of Piezodorus hybneri, Leptocorisa sp., 
Leptocorisa acuta, Riptortus sp. (Hemiptera: 
Alydididae), Amblypelta lutescens papuensis, 
Dasynus piperis, Mictis prof ana (Hemiptera: 
Coreidae), Nezara viridula (Hemiptera: Pentato- 
midae), an unidentified coreid bug and an uni- 
dentified moth. 

Use in biocontrol. O. utetheisae is almost cer- 
tainly the species introduced into Kenya and 
Tanzania as Ooencyrtus sp. for the control of 
Pseudotheraptus wayi (Hemiptera: Coreidae) 
(see Greathead, 1971). This may also have been 
the species introduced, under the name of Ooen- 
cyrtus malayensis, into the Solomon Islands in 
1937-1938 for the control of Amblypelta 
cocophaga (Hemiptera: Coreidae) (Phillips, 
1941) and the United States (Massachussets) in 
1981 for the control of Anasa tristis (Hemiptera: 
Coreidae) and Nezara viridula (Hemiptera: Pen- 
tatomidae) (Coulson et al. 1988). 

Distribution. Sub-saharan Africa, India, 
Nepal, Thailand, P.R. China, Philippines, 
Malaysia, Indonesia, Papua New Guinea. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



83 



Material examined. Type material. Holotype 
9 of Ooencyrtus patriciae Subba Rao, NIGE- 
RIA: Kaduna State, Yankara, ex Clavigralla 
tomentosicollis (P.C. Matteson). 

Non-type material. INDIA: 1$, Uttar 
Pradesh, Dehra Dun, x.1979 (Z. Boucek); 1$, 
Karnataka, Bangalore, 19-23. ix. 1979 (J.S. 
Noyes); 49, Tamil Nadu, Coimbatore, 
25. ix-1. x.1979 (J.S. Noyes); 29, Kerala, Wala- 
yar Forest, 26.xi-l. x.1979 (J.S. Noyes); 29, 
Periyar A. Sane. 5-15.X.1979 (J.S. Noyes); 2$, 
Assam, Tinsukia, on eggs of Hemiptera, HE 
21669, 12.viii.1990 (F. Alam); 59, Assam, Tin- 
sukia, ex eggs of Leptocorisa, HE 22252, 
28.vii.1991 (Md. Farouk); 39, Assam, Tinsukia, 
29.vii.1991 (Md. Farouk), 19, Assam, Tinsukia, 
HE A20982, 16. vi. 1989; 29, Assam, Tinsukia, ex 
eggs Nezara viridula, HE 22252, 7.viii. 1991 (Md. 
Farouk); 39, ?, CLE. A17567; NEPAL: 19, 
Kakani, 2070m, x.1983 (M.G. Allen); 29, Chit- 
wan, Inner Terai, 150m, x.1983 (K.K. Gurung); 
159, 2d", Maharashtra, Dapoli, ex egg bug on 
snake gourd, CIE 19243, 1987; THAILAND: 
19, Nan Pr. Nan, edge of rice paddy, BM 
1986-78, 31.viii.1985 (M.J.E. Reacher-Huber); 
29, Huai Kha Khaeng, ii.1986 (M.G. Allen); 
19, Chang Kiang, Doi Suthop, ii.1985 (D. Jack- 
son); 149, 60", Ratismoa Prov. Uthang Dist., ex 
eggs of Piezodorus hybneri & Nezara viridula, 
19-20. viii. 1989 (M. Kogan) [identified as Ooen- 
cyrtus malayensis Ferriere by B.R. Subba Rao]; 
P.R. CHINA: 49 , Canton (Guangdong), ex eggs 
of a coreid bug (R.B. Falkenstein); 19, Hainan, 
Dang Xiang, 7.V.1983 (Z. Boucek); PHILIP- 
PINES: 249, 15a", Laguna, Los Banos, 150 ft. 
ex Leptocorisa, v. 1930 (L.B. Uichanco); 
MALAYSIA: Setapak, 59, 2o\ ex eggs of Lep- 
tocorisa sp., 17.xii.l922(G.H. Corbett & B.A.R. 
Gater) [syntypes of Ooencyrtus malayensis]; 19, 
Sumger Lua, ex eggs of Leptocorisa acuta Thun- 
berg, 20.xi.1928 (G.H. Corbett) [syntype of 
Ooencyrtus malayensis]; 19, Genting Highlands, 
vii.1981 (R.I. Vane Wright); 49, Sarawak, 4th 
div. Gn. Mulu., RGS Exp. 17.ix-23.x.l977 (D. 
Hollis); 39, Sarawak, Long Lama, 13. ii. 1987 
(A.T. Finamore & C. Boxfield); 19, Sabah, 
Kinabalu NP, Poring Hot Springs, 12. v. 1987 (A. 
Smetana); 19, Sabah, Kinabalu NP, Poring Hot 
Springs, 900m, 14.V.1987 (A. Smetana); 1$, 
Sabah, Kinabalu NP, Poring Hot Springs, 500m, 
30.viii.1988 (A. Smetana); 29, Sabah, Danum 
Valley, xi.1986 (P. Eggleton); INDONESIA, 
69, Banka Is. ex Dasynus piperis China, 1930 
(J. van der Vecht); 39, 2d\ Sumatra, Pematang 
Siantar, ex eggs of a moth, 30. ix. 1931 (R.I. Nel); 
19, Java, Bogor, viii. 1983 (Hull Univ. Exp.); 
49, Java, Krakatau, various localities and dates 



ix.1984 (S.G. Compton); 29, Sulawesi Tengah, 
Nr. Morowali, Ranu River Area, Lowland rain 
forest, hi. 1980 (M.J.D. Brendell); 929, 
Sulawesi, Utara, Dumoga-Bone NP, Toraut, 
iv.1985 (J.S. Noyes); PAPUA NEW GUINEA: 
69, Morobe Prov. Markham Valley, ex eggs of 
Riptortus sp., CIE A12822, 23.xii.1980 (G 
Youma); 19, Areme, Musgrave R. 9.xii. 1982 
(Z. Boucek); l9,Mt. Hagen, 1600m, 17.xii.1982 
(Z. Boucek); 109, 3d", Laloki Res. St. ex eggs 
of Amblypelta lutescens papuensis, 24. viii. 1989 
(F.M. Don). Material in BMNH, ZISP, QMB, 
MZB, PPRI, CNC, USNM. 

Extralimital material. SENEGAL: 39, Bam- 
bey, ex eggs of coreid, 21.x. 1943 (J. Risbec); 
P.R.BENIN: 49, 5d\ Cotonou, ex egg of Clavi- 
gralla shadabi, 1992 (H. Dreyer); 1$, Id", Coto- 
nou, ex egg Riptortus dentipes 1992 (H. Dreyer); 
NIGERIA: 29, Id", Ibadan, ex eggs Anoplocne- 
mis curvipes, CIE A8523, 1975 (Ochiah); 
MALAWI: 59, ree'd 10.xii.1992, Lab reared ex 
eggs of Bathycoelia bequarti, Rec'd from D.A. 
Ironside /93-001, HE 22730; TANZANIA: 39, 
60", Zanzibar, ex eggs of Pseudotheraptus wayi, 
1959, CIE Coll. 16944 (B.H. Hyde-Wyatt). 
Material in BMNH. 

Comments. Prinsloo (1987) treated Aenasioidea 
demodoci both as a synonym of utetheisae (his 
page 9) and as a valid species (his page 15). This 
cannot be resolved until a lectotype is designated 
for demodoci. Clearly it would be better to 
designate the lectotype from the series belonging 
to the valid species as interpreted by Prinsloo on 
page 15 of his paper and therefore we are not 
including the name here as a synonym of utethe- 
isae. 

This species can be confused easily with papil- 
ionis as demonstrated by the inclusion of several 
specimens of utetheisae in the syntypic series of 
malayensis by Ferriere. Females of utetheisae can 
be separated relaibaly from papilionis by the 
dark brown ovipositor sheaths whereas in papil- 
ionis the sheaths are yellow or orange. Otherwise 
the flagellar segments of utetheisae are relatively 
longer and the structure of the ovipositor and 
hypopygium is slightly different (compare Figs 
317-319 with 329 and 331, Figs 324-326 with 
336). 



84 



D. -W.HUANG AND J.S. NOYES 



EXCLUDED SPECIES 



Ooencyrtus johnsoni (Howard) 

Encyrtus johnsoni Howard, 1898: 18. Syntypes 
$Cf , USA (USNM, not examined. 

Ooencyrtus johnsoni (Howard); Ashmead, 1900: 
382. 

Comments. This species was introduced into 
Hawaii in 1940 for the control of Murgantia 
hislrionica (Hemiptera: Pentatomidae). 

Although Fullaway (1947) reported that the spe- 
cies was well established in 1946 it has not been 
reported in Hawaii since then (Beardsley, 1976). 
We have not seen any specimens of this species 
from Hawaii and therefore it is not included here 
in the treatment of species. 

Ooencyrtus trinidadensis Crawford 

Ooencyrtus trinidadensis Crawford, 1913: 347. 
Lectotype $, Trinidad (USNM, not exam- 
ined). 

Comments. Introduced into Hawaii in 1962 for 
the control of Nezara viridula (Hemiptera: Pen- 
tatomidae) but apparently did not become estab- 
lished (Davis, 1964; Waterhouse & Norris, 
1987). 

The species is very close to, and may be 
synonymous with, Ooencyrtus johnsoni 
(Howard). 



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Prinsloo, G.L. 1987. A revision of the genus Ooencyrtus 
Ashmead (Hymenoptera: Encyrtidae) in sub-saharan Africa. 
Entomology Memoir, Department of Agriculture and Water 
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Prinsloo, G.L. & Annecke, D.P. 1978a. Two new species of 
Ooencyrtus (Hym. Encyrtidae) parasitic in Othreis fullonia 
(Clerck) (Lep. Noctuidae) from New Caledonia and Western 
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Technique Outre-Mer (Serie Biologie) 13(1): 41-44. 

Prota, R. 1966. Contributi alia conoscenza dell'entomofauna 
della Quercia da sughero Quercus suber L.) - V - Osser- 
vazioni condotte in Sardegna su Ooencyrtus kuwanai (How.) 
(Hymenoptera Encyrtidae) nuovo pe la fauna italiana. Stazi- 
one Sperimentale de; Sughero. Tempio Pausania, Memoria 
17: 3-26. 

Rahim, A. & Hashmi, A. A. 1984. Biological control of Pyrilla 
perpusilla in Sind, Pakistan. International Pest Control 26(5): 
124-126. 

Rahim, A., Hashmi, A. & Khan, N.A. 1991. Effects of tem- 
perature and relative humidity on longevity and develop- 
ment of Ooencyrtus papilionis Ashmead (Hymenoptera: 
Eulophidae), a parasite of the sugarcane pest Pyrilla perpu- 
silla Walker (Homoptera: Cicadellidae). Environmental 
Entomology 20(3): 774-775. 

Rajak, R.L., Pawar, A.D., Misra, M.P., Prasad, Varma, A. & 
Singh, G.P. 1987. Sugarcane pyrilla epidemics 1985 - a case 
study. Plant Protection Bulletin 39: 1-9. 

Risbec, J. 1951. 1. Les Chalcidoides de l'Afrique occidentale 
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1954. Chalcidoides et Proctotrupoides de l'Afrique occi- 
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1958. Contributions a la connaissance des Hymenopteres 

Chalcidoides et Proctotrupoides de l'Afrique Noire. Annates 
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Romanova, V.P. 1953. Egg parasites of harmful Pentatomidae 
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Shafee, S.A., Alam, M. & Agarwal, M.M. 1975. Taxonomic 
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Stouthamer, R. 1990. Evidence for microbe-mediated parthe- 
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Stouthamer, R.. Luck, R.F. & Hamilton, W.D. 1990. Antibiot- 
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National Academy of Sciences USA 87(7): 2424-2427. 

Subba Rao, B.R. 1979. Taxonomic studies on some encyrtid 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



87 



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1981. Description of Ooencyrtus patriciae n.sp. from 

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new species of parasitic Ooencyrtus Ashmead, 1900 (Hym., 
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Vassiliev, I. 1904. Ueber eine neue, bei den Vertretern der 
Gattung Telenomus parasitierende Encyrtus-Art 



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Waterston, J. 1915. Ooencyrtus pacificus a new egg parasite 
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Ooencyrtus kuwanai. Annals of the Entomological Society of 
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longevity of gypsy moth (Lepidoptera: Lymantriidac) egg 
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Density, size and mortality of egg masses in New Jersey 
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1960b. Environmental determination of sex in a partheno- 
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Yadav, R.P. & Chaudhary, J. P. 1984. Laboratory studies on 
the biology of Ooencyrtus papilionis Ashmead 
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Entomological Research 8(2): 162-166. 

1985. Upper limit of temperature for the development of 

Ooencyrtus papilionis Ashmead within its host eggs. Current 
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Pest Management 28(2): 107-1 14. 



88 



D. -W.HUANG AND J.S. NOYES 




Figs 1-8 O. guamensis- 1, antenna including reverse side of clava, 9; 2, scutellum, 9; 3, right fore wing, upper 
aspect, 9; 4, antenna with clava in side aspect showing oblique truncation, 9; 5, hypopygium; 6, antenna, cf; 7, 
foretibia showing inner, apical scaly area, cf ; 8, genitalia cf . 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



89 




m'/dU- 



iW/t 



13 



1 //// 




Figs 9-15 O. acca - 9, antenna, 9; 10, left mandible; 11, right forewing, upper surface, $; 12, scutellum, §; 13, 
right forewing, upper surface, $; 14, ovipositor; 15, hypopygium. 



90 



D. -W.HUANG AND J.S. NOYES 




Figs 16-21 O. phongi - 16, right antenna, outer aspect including reverse side of clava, $ ; 17, head, facial view, 
$; 18, left mandible, 9; 19, right forewing, upper surface, $; 20, genitalia, d"; 21, antenna, O". 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



91 




Figs 22-27 O. acus - 22, antenna including reverse side of clava, $; 23, right forewing, upper surface, 9; 24, 
ovipositor; 25, hypopygium; 26, antenna, cf ; 27, genitalia, cf . 



92 



D. -W.HUANG AND J.S. NOYES 




Figs 28-33 O. caurus - 28, antenna including reverse side of clava, 9 ; 29, right forewing, upper surface, 9 1 30, 
ovipositor; 31, hypopygium; 32, antenna, cf ; 33, genitalia, cf. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



93 







jf/'; /JAy, ' fir 



si's; s ' 



>£ 





Figs 34-38 O. lucens - 34, antenna including reverse side of clava,; $ ; 35, right forewing, upper surface, $; 36, 
hypopygium; O. minerva - 37, antenna, $; 38, right forewing, upper surface, $. 



94 



D.-W. HUANG AND J.S. NOYES 




Figs 39-46 O. icarus - 39, hypopygium; 40, right forewing, upper surface, $ ; 41 , left mandible. O. adonis - 42, 
antenna, 9; 43, head, facial aspect, §; 44, right forewing, upper surface, $. O. aeneas - 45, head, facial aspect, 
9; 46, left mandible. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



95 




Figs 47-53 O. aeneas - 47, antenna, 9; 48, right forewing, upper surface, $; 49, ovipositor; 50, hypopygium. O. 
kuvanae- 51, antenna, Q; 52, hypopygium; 53, right forewing, upper surface, $. 



96 



D. -W.HUANG AND J.S. NOYES 




Figs 54-60 O. dis - 54, antenna, $; 55, right forewing, upper surface, 9'» 56, head, facial aspect, $; 57, 
hypopygium; 58, ovipositor. O. hercle - 59, antenna, 9; 60, right forewing, upper surface, 9- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



97 




Figs 61-66 O. hercle- 61, hypopygium; 62, head, facial aspect, 9; 63, antenna, cf. O. corbetti- 64, antenna 
clava slightly collapsed, $; 65, right forewing, upper surface, 9; 66, ovipositor. 



98 



D. -W.HUANG AND J.S. NOYES 




y 



'"/'s/ 

y^^/ 



/i>M 





Figs 67-75 O. podontiae - 67 , antenna, 9 paralectotype,; 68, antenna, 9 non-type,; 69, left mandible, 9; 70, 
right forewing, upper surface, 9; 71, scutellum, 9; 72, ovipositor; 73, hypopygium. O. pindarus - 74, head, 
facial aspect, 9; 75, left mandible. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



99 




Figs 76-82 O. pindarus - 76, antenna, 9; 77, right forewing, upper surface; $; 78, ovipositor; 79, genitalia, C? ; 
80, scutellum, 9; 81, hypopygium; 82, right antennae, outer aspect, cf- 



100 



D. -W.HUANG AND J.S. NOYES 




Figs 83-88 O. plautus - 83, antenna, $; 84, right forewing base, upper surface, 9; 85, head, facial aspect, $; 86, 
ovipositor; 87, genitalia, cf; 88, antenna, cT. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



101 




Figs 89-95 O. daphne - 89, antenna, 9; 90, right forewing base, upper surface, $; 91, left mandible; 92, 
hypopygium; 93, ovipositor, 9- O. flavipes - 94, 95, head, facial aspect, $. 



102 



D. -W.HUANG AND J.S. NOYES 




Figs 96-101 O. flavipes - 96, 97, antenna, <j>; 98, 99, right forewing base, upper surface, 9; 100, ovipositor; 101, 
hypopygium. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



103 




Figs 102-107 O. flavipes - 102, hypopygium. O. dione - 103, right forewing base, upper surlace. Q; 104, antenna 
including reverse side of clava which is slightly collapsed, 9: 105, scutellum, $. O. dry as - 106, scutellum, Q; 
107, right forewing base, upper surface, 9- 



104 



D. -W.HUANG AND J.S. NOYES 




Figs 108-115 O. ceres - 108, antenna, $; 109, scutellum, §; 110, right forewing base, upper surface, 9; HI. 
head, facial aspect, 9; U2, hypopygium. O. ition - 113, antenna including reverse side of clava, 9; 114, left 
mandible; 115, right forewing base, upper surface, 9- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



105 




,' 



''W/'f^s&j' 




Figs 116-123 O. ilion - 116, ovipositor. O. iris- 117, head, facial aspect, 9; 118, hypopygium; 119, right 
forewing base, upper surface, 9- O. javanicus - 120, antenna, 9; 121, left mandible; 122, ovipositor; 123, right 
forewing base, upper surface, 9- 



106 



D. -W.HUANG AND J.S. NOYES 




Figs 124-130 O. javanicus- 124, antenna, d". O. lulus- 125, 126, antenna, variants, $ ; 127, head, facial aspect, 9 ; 128, 
right forewing, upper surface, $ . O. ixion - 129, antenna, 9 ; 130, right forewing base, upper surface, $ . 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



107 








W j/ 






Figs 131-137 O. ixion- 131, ovipositor; 132, head, facial aspect, 9- O. elissa- 133, head, facial aspect, $; 134, 
antenna, 9; 135, right forewing base, upper surface, $; 136, hypopygium; 137, left mandible. 



108 



D. -W.HUANG AND J.S. NOYES 




Figs 138-144 O. egeria - 138, head, facial aspect, $; 139, leaft mandible; 140, right forewing base, upper surface, 
9- O. erebus - 141, 142, antenna, variants, $; 143, left mandible; 144, right forewing base, upper surface, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



109 




Figs 145-153 O. erebus- 145, ovipositor; 146, scutellum, $; 147, right forewing base, upper surface, $. O. 
endymion - 148, antenna, $; 149, hypopygium; 150, 151, left mandible, variants; 152, ovipositor; 153, right 
forewing, upper surface, 9- 



no 



D. -W.HUANG AND J.S. NOYES 




Figs 154-161 O. hymen - 154, antenna, $; 155, left mandible; 156, right forewing base, upper surface; 157, 
ovipositor; 158, head, facial aspect, $. O. belus - 159, head, facial aspect, J; 160, right antenna, outer apsect, 
$; 161, right forewing base, upper surface, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



111 




Figs 162-168 O. belus - 162, ovipositor; 163, hypopygium; 164, antenna, cf ; 165, genitalia, cf • O. bacchus 
hypopygium; 167, antenna, $; 168, right forewing base, upper surface, $. 



166, 



112 



D. -W.HUANG AND J.S. NOYES 




Figs 169-174 O. pacificus - 169, antenna, 9; 170, right forewing base, upper surface, $; 171, ovipositor; 172, 
head, facial aspect, $; 173, left mandible; 174, antenna, cf . 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



113 




Figs 175-182 O. larvarum - 175, antenna, $; 176, left mandible; 177, hypopygium; 178, head, facial aspect, a"; 
179, ovipositor; 180, antenna, $ . O. pallidipes - 181, antenna, $ ; 182, right forewing base, upper surface, 9 • 



114 



D. -W.HUANG AND J.S. NOYES 




Figs 183-189 O. pallidipes - 183, head, facial aspect, 9; 184, right forewing base, upper surface, Q\ 185, 186, 
hypopygium, variants, $. O. pilosus - 187, antenna, 9; 188, left mandible, $; 189, right forewing, upper 
surface, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



115 




Figs 190-195 O. valcanus- 190, antenna including reverse of clava, 9; 191, hypopygium; 192, right forewing, 
upper surface, 9- O. utania - 193, antenna including reverse of clava, 9; 194, hypopygium; 195, right forewing 
base, upper surface, 9- 



116 



D. -W.HUANG AND J.S. NOYES 




Figs 196-205 O. vertumnus - 196, antenna including reverse of clava, clava slightly collapsed,; $?> 197, head, 
facial aspect, $; 198, right forewing base, upper surface, $; 199, ovipositor; 200, hypopygium. O. vesta- 201, 
head, facial aspect, 9; 202, antenna, $. O. boreas- 203, hypopygium; 204, left mandible; 205, ovipositor. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



117 




Figs 206-211 O. boreas - 206, antenna, $; 207, right forewing base, upper surface; $>. O. clio - 208, antenna, £; 
209, right forewing base, upper surface, 9; 210, head, facial aspect, $ ; 211, hypopygium. 



118 



D. -W.HUANG AND J.S. NOYES 








Figs 212-220 O. circe- 212, antenna, §; 213, left mandible; 214, right forewing base, upper surface, 9; 215, 216, 
head, facial aspect, variants, $; 217, hypopygium. O. clotho - 218, hypopygium; 219, head, facial aspect, 9; 
220, ovipositor. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



119 




Figs 221-227 O. cybele- 221, antenna, $; 222, right forewing base, upper surface, 9; 223, hypopygium; 224, 
head, facial aspect, $; 225, ovipositor. O. ooii- 226, ovipositor. O. crassulus - 227 , antenna, 9- 



120 



D. -W.HUANG AND J.S. NOYES 










//'/'/ 

?$■&£' 

'/?'-'- 



mmmm 



/ 






W/W'Wr'. 






232 



mMw- 







Figs 228-232 O. lucina - 228, right antenna, outer aspect, 9; 229, right forewing base, upper surface, £; 230, 
ovipositor; 231, antenna, d\ O. macula - 232, right forewing base, upper surface, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



121 




Figs 233-241 O. macula - 233, antenna, 9; 234, head, facial aspect, 9; 235, hypopygium; 236, ovipositor; 237, 
genitalia, cT; 238, antenna, d". O. hera- 239, head, facial aspect, 9; 240, left mandible, 9; 241, hypopygium. 



122 



D. -W.HUANG AND J.S. NOYES 




Figs 242-249 O. hera - 242, antenna, §; 243, right forewing base, upper surface, $; 244, ovipositor; 245, 
antenna, cf- O. manii- 246, scutellum, $; 247, head, facial aspect, $; 248, left mandible; 249, right forewing 
base, upper surface, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



123 




Figs 250-258 O. manii - 250, 251, antennae, variants, $; 252, ovipositor; 253, 254, hypopygium, variants, $. O. 
midas- 255, head, facial aspect, 9; 256, antenna, Q; 257, right forewing base, upper surface, ty; 258, 
ovipositor. 



124 



D. -W.HUANG AND J.S. NOYES 




Figs 259-265 O. midas - 259, hypopygium. O. mars - 260, head, facial aspect, 9 ; 261, right forewing base, upper 
surface, $; 262, antenna, $; 263, hypopygium; 264, ovipositor. O. neptunus - 265, right forewing base, upper 
surface, 9- 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



125 




Figs 266-274 O. neptunus - 266, ovipositor, $; 267, scutellum, 9; 268, left mandible; 269, hypopygium. O. 
libitina - 270, antenna, $; 271, right forewing base, upper surface, $; 272, ovipositor. O. lyaeus-273, 
hypopygium; 274, ovipositor. 



126 



D. -W.HUANG AND J.S. NOYES 




Figs 275-281 O. lyaeus - 275, antenna including reverse side of clava, $; 276, right forewing base, upper aspect, 
9- O lupercus - 211 , antenna showing reverse side of clava, $ ; 278, right forewing, upper surface, 9 ! 279, 
ovipositor; 280, head, facial view, 9; 281, hypopygium. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



127 




Figs 282-291 O. maenas - 282, antenna including reverse side of clava, 9; 283, left mandible; 284, ovipositor; 
285, hypopygium. O. segestes - 286, head, facial aspect, 9; 287, antenna, $; 288, 289, scutellum, variants, 9; 
290, left mandible; 291, right forewing, upper surface, brachypterous form, variant, 9- 



128 



D. -W.HUANG AND J.S. NOYES 






Figs 292-298 O. segestes - 292, right forewing, upper surface, brachypterous form, variant, 9; 293, right 
forewing base, upper surface, macropterous form, $; 294, ovipositor; 295, antenna, d". O ferrierei - 296, head, 
facial view, 9; 297, antenna, J; 298, hypopygium. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



129 




Figs 299-307 O. ferrierei - 299, ovipositor. O. leander - 300, antenna, 9 1 301 , right forewing base, upper surface, 
$; 302, ovipositor; 303, laft mandible. O. musa - 304, antenna, $; 305, scutellum, $; 306, hypopygium; 307, 
ovipositor. 



130 



D. -W.HUANG AND J.S. NOYES 




Figs 308-316 O. telenomicida - 308, antenna, 9 ! 309, scutellum, $ ; 310, right forewing base, upper surface; 311, 
ovipositor 312, head, facial aspect, 9; 313, hypopygium. O. papilionis - 314, 315, head, facial aspect, variants, 
$; 316, scutellum, $. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



131 




7/ 7 M# 






Figs 317-323 0. papilionis - 317-319, antenna, variants, 9 \ 320, 321, right forewing base, upper surface, 
variants, $; 322, 323, ovipositor, variants. 



132 



D. -W.HUANG AND J.S. NOYES 








Figs 324-330 O. papilionis - 324-326, hypopygium, variants, $; 327, head, facial aspect, cf ; 328, right antenna, 
outer aspect, cT. O. utetheisae - 329, antenna, $ ; 330, right forewing base, upper surface. 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



133 




Figs 331-337 O. utetheisae - 331, antenna, 9; 332, right forewing base, upper surface, $ ; 333, 334, left 
mandible, variants; 335, ovipositor; 336, hypopygium; 337, antenna, cf. 



134 



D. -W.HUANG AND J.S. NOYES 



INDEX 



Synonyms are in italics: main citations in bold. 



Abelmoschus 21 , 22 
acastus (Ooencyrtus) 79 
acca (Ooencyrtus) 11, 17, 18, 89 
Achaea 6, 22 
Acrocercops 5, 62 
acus (Ooencyrtus) 5, 11, 19, 91 
acuta (Leptocorisa) 4, 80, 82. 83 
acuta (Turnaca) 6, 65 
acutangulatus (Gonocerus) 5, 78 
Adelencyrtus 31 

adonis (Ooencyrtus) 11. 24. 25, 26. 94 
aegeus (Papilio) 6, 79, 80 
aegyptius (Ischiodon) 4. 16, 17 
Aelia 5, 78 
Aenasioidea 82, 83 

aeneas (Ooencyrtus) 12, 25, 26, 94, 95 
afer (Ooencyrtus) 27 
affinis (Podontia) 30 
agamemnon (Papilio) 6. 79, 80 
Ageniaspis 67 
Agiommatus 26 
Agonoxena 5, 81 
Agrilus 51 
Allograpta 4, 16 

Amblypelta 4, 8, 21, 22. 38, 79. 82, 83 
americana (Malacosoma) 6, 27 
Amorpha 7, 78 
Anagyropsis 81 
Anasa 8, 79. 82 
anasae (Ooencyrtus) 3 . 
Anastatus 2, 26, 27 
Anoplocnemis 4, 82, 83 
Apanteles 2, 9, 27 
Aphis 16, 17 
Ariadne 6, 79, 80 
ariadne (Ariadne) 6, 79, 80 
Aroa 6, 79, 80 
Arrhenophagus 51 
arundinacea (Imperata) 75 
Asarkira 16, 17 
Ascotis 8 

atlas (Attacus) 6, 26 
atomosa (Exelastis) 6, 16 
attaci (Agiommatus) 26 
Attacus 6, 26 
auritus (Paragus) 4, 16 
austanti (Amorpha) 7, 78 
australiensis (Ooencyrtus, 
Tetracnemella) 9 

bacchus (Ooencyrtus) 13, 37, 39, 43, 49, 

50, 111 
Bathycoelia 5, 83 

belus (Ooencyrtus) 13, 47, 48, 110, 111 
bequarti (Bathycoelia) 5, 83 
bifasciatus (Anastatus) 2 
bolina (Hypolimnas) 6, 79, 80 
Borassus 75 

borbonicus (Paragus) 16, 17 
boreas (Ooencyrtus) 6, 14, 57, 116, 117 
Brachynema 5, 78 
Brachyplatys 5, 50 
brazai (Orianos) 51 



Caligo6, 51,52 

Callosamia 7, 27 

Calpodes 8 

calpodicus (Ooencyrtus) 8 

catappa (Terminalia) 62 

caurus (Ooencyrtus) 4, 11. 21. 24, 92 

celerio (Hippotion) 7, 63 

Cephanodes 7, 79, 80 

ceres (Ooencyrtus) 5. 12, 23, 37. 38, 39, 

43, 49, 53, 104 
charitonius (Hcliconius) 6. 79, 80 
Chilasa 79, 80 
Chilo 5. 79, 80 
Chromus 81 

chrysippus (Danaus) 5. 80 
chrysorrhoca (Euproctis) 6, 27 
circe (Ooencyrtus) 14. 59, 60, 61, 118 
citri (Papilio) 6, 32 
citrifolia (Morinda) 81 
Clavigralla 4, 5, 82. 83 
clio (Ooencyrtus) 14, 58. 59. 60. 61, 117 
clisiocampae (Encyrtus, Ooencyrtus) 8 
Clostera 6, 64 

clotho (Ooencyrtus) 14, 60. 61. 118 
cochereaui (Ooencyrtus) 79. 80 
cocophaga (Amblypelta) 4, 8, 38. 79, 82 
Colocasia 80 

columbi (Mirsyrpophagus) 81 
cometaris (Aroa) 6, 79, 80 
compositae (Dactynotus) 16. 17 
Conopomorpha 5. 8. 62, 97 
corbetti (Ooencyrtus) 4, 12, 29. 30. 31 
core (Euploea) 5, 79, 80 
craccivora (Aphis) 16, 17 
cramerella (Conopomorpha) 5, 8, 62 
crassulus (Ooencyrtus) 6. 7. 14, 63. 63, 

81 
Cratoplatys5,37 
creona (Litomastix, Ooencyrtus, 

Pseudolitomastix) 9 
Cricula 7, 35, 40 
cupreata (Clostera) 6, 64 
curvipes (Anoplocnemis) 4. 82, 83 
cybele (Ooencyrtus) 4, 14, 61, 62, 119 

Dactynotus 16, 17 

Danaus 5, 80 

daphne (Ooencyrtus) 12, 33, 34, 101 

Dasynus5,24, 82, 83 

definata (Hemerocampa) 6, 27 

Delias 6, 50, 51 

demodoci (Aenasioidea, Ooencyrtus) 83 

demoleus (Papilio) 6, 32, 79. 80 

Dendrolimus 6, 27, 46 

dentipes (Riptortus) 4, 82 

destructor (Echthrodryinus, Ooencyrtus) 

8 
Dideopsis 4 
Dimocarpus 19 

dione (Ooencyrtus) 7. 12, 35. 103 
dipterae (Ooencyrtus) 43 
dis (Ooencyrtus) 12, 27, 28, 96 
dispar (Lymantria) 2, 6. 7, 8, 9, 27 



disparis (Anastatus) 27 
distatus (Ooencyrtus) 54 
Dolycoris 5. 78 
dryas (Ooencyrtus) 12, 36, 103 

Echthrodryinus 8. 9. 19. 20 

Ectopiognatha 9. 77 

egeria (Ooencyrtus) 13, 44, 108 

elaezia (Cricula) 7, 35 

elissa (Ooencyrtus) 13, 37, 39, 42, 43. 

44. 49, 53, 62, 77, 107 
elongata (Clavigralla) 5, 82 
endymion (Ooencyrtus) 6, 7, 13, 46. 47, 

109 
Eocanthecona 5. 41 
Epipyrops 20 
Epiricania 5, 20 
eras (Gnathothlibas) 7. 81 
erebus (Ooencyrtus) 13, 44, 45. 108, 109 
Eriogyna 7, 27 
Erionota6, 8, 51,52, 79 
erionotae (Ooencyrtus) 51, 62 
erotus (Gnathothlibas) 7, 81 
esculenta (Colocasia) 80 
ethlius 8 

Euploea 5. 79, 80 
Euproctis 6, 27 
Eurema 6. 50, 51 
Eurygaster 5, 8, 78 
Exelastis 6. 16 
exotica (Allograpta) 4, 16 

fecundus (Ooencyrtus) 2, 3. 7, 8, 46 
ferrierei (Ooencyrtus) 4, 15. 42, 76. 78. 

128, 129 
flavofasciatus (Schedius) 78 
flavipes (Ooencyrtus, Xesmatia) 9, 12. 

33.34. 101. 102. 103 
fullonia (Othreis) 6. 8, 63, 79. 80 
fumida (Lymantria) 6, 27 
furcellata (Eocanthecona) 5. 41 

garouae (Ooencyrtus. Schedius) 16. 17 
germarii (Brachynema) 5. 78 
glaucippe (Hebomoia) 6, 66 
globulifera (Acrocercops) 5, 62 
Gnathothlibas 7 
goarini (Psyllaephagus) 16, 17 
gonoceri (Ooencyrtus) 79 
Gonocerus 5, 78 

guamensis (Ooencyrtus) 4. 6, 11. 16, 17, 
18,88 

Hadart 52 

Hasora 6, 79, 80 

Hebomoia 6, 66 

Helegonatopus 9, 64 

helena (Troides) 6. 79. 80 

helenus (Papilio) 6, 80 

Heliconius 6, 79, 80 

Hemerocampa 6, 8, 27 

Hemileuca 7, 8, 27 

hera (Ooencyrtus) 6, 14. 66. 81, 121, 122 



REVISION OF INDO-PACIFIC SPECIES OF OOENCYRTUS 



135 



hcrcle (Ooencyrtus) 5, 12. 28. 29. 96, 97 

Hippotion 7, 63 

histrionica 8. 83 

hybneri (Piezodorus) 4, 82. 83 

hylas (Cephanodes) 7. 79. 80 

hymen (Ooencyrtus) 13, 47, 48, 1 10 

Hypolimnas 6, 79, 80 

icarus (Ooencyrtus) 5, 11, 23, 24, 94 
ilion (Ooencyrtus) 4, 12, 38, 41, 104, 105 
Imperata 75 
indica (Mangifcra) 62 
integriccps (Eurygaster) 5, 8. 78 
iris (Ooencyrtus) 12, 24, 37, 39, 43, 49 
Ischiodon 4, 16, 17 
isonoma (Acroccrcops) 63 
iulus (Ooencyrtus) 5, 13. 38. 40. 41, 106 
ixion (Ooencyrtus) 7, 13, 41, 42, 106, 
107 

jaculus (Mirpcrus) 4, 82 
Janata (Achaea) 6, 22 
javanica (Tcssaratoma) 5,19 
javanicus (Ooencyrtus, Schcdius) 5, 7, 

13.40,41, 105, 106 
johnsoni (Encyrtus, Ooencyrtus) 3, 8, 

83,84 
julii (Tanaccia) 6, 80 
juniperi (Gonocerus) 5, 78 
Junonia 6, 79 

Kallima6, 79, 80 
kalshovenac (Dasynus) 5, 24 
kikuchii (Dcndrolimus) 6, 46 
kuvanae (Ooencyrtus, Schcdius) 2. 3, 4, 

6,7,8. 10, 12,26.27,95 
kuwanae (Ooencyrtus) 26 
kuwanai (Ooencyrtus) 26 

larvarum (Ooencyrtus, 

Paracopidosomopsis) 6. 13, SO, 66. 1 13 
leander (Ooencyrtus) 9. 15, 76. 77. 129 
lemonias (Junonia) 6. 79, 80 
Leptocorisa 4, 80, 82,83 
leucocerus (Ooencyrtus) 79, 80 
leucostigma (Hemerocampa) 6, 8, 27 
libitina (Ooencyrtus) 15, 56. 68. 69. 71. 

72, 74, 125 
lichenina (Stauropus) 6, 58 
limniacc (Tirumala) 5, 79, 80 
Litomastix 9 
longan (Dimocarpus) 19 
lucens (Ooencyrtus) 5. 6, 11. 20. 21, 22. 

24, 93, 120 
lucina (Ooencyrtus) 6, 14, 64. 120 
lupercus (Ooencyrtus) 15, 71, 72, 73, 74, 

126 
lutescens (Amblypelta) 4, 21, 22, 82, 83 
lyaeus (Ooencyrtus) 15, 71, 72, 73, 74, 

125, 126 
Lymantria 2, 6. 7. 8. 9. 27 

macula (Ooencyrtus) 6, 65, 120, 121 
maenas (Ooencyrtus) 15, 57, 71, 72. 73. 

74, 127 
maia (Hemileuca) 7, 27 
major (Ooencyrtus) 26 
major (Ectopiognatha, Ooencyrtus) 77 
Malacosoma 6, 27 
malayensis (Ooencyrtus) 8, 79, 80, 82, 83 



Mangifcra 62 

manihoti (Abelmoschus) 21, 22 

manihoti (Phcnacoccus) 17 

manii (Ooencyrtus) 3, 4, 5, 15, 67, 69, 

79. 122, 123 
mars (Ooencyrtus) 15, 69. 124 
mclanoleuca (Epipvrops. Epiricania) 5. 

20 
mclanoscelus (Apanteles) 2. 9. 27 
memnon (Caligo) 6, 51. 52 
memnon (Papilio) 6, 79. so 
menciana (Rondotia) 5. 29 
menzcli (Anastatus) 26 
Mctaphycus 64 
Microterys 9 
Mictis 5, 82 
midas(Oocncvrtus) 5. 15.67.68.69.71. 

72, 123, 124 
mimus (Ooencyrtus) 54 
minerva (Ooencyrtus) 1 1, 23. 93 
minor (Ectopiognatha. Ooencyrtus) 9. 

77 
Mirpcrus 4, 82 
Mirsyrpophagus 81 
Morinda 81 
Murgantia 8. 83 

nuisa (Ooencyrtus) 15. 75. 77, 78. 129 
Mythimna 6, 16 

neptunus (Ooencyrtus) 15.70. 124. 125 

ncustna (Malacosoma) 6. 27 
Nezara 5, 8, 80. 82. 83. 84 
nezarac (Ooencyrtus) 3. 8 

nigerrimus (Ooencyrtus) 3. 8 
Nygmia 6, 27 

oliviac (Hemileuca) 7. 8. 27 

ooii (Ooencyrtus) 5. 8. 14. 62, 63, 1 19 

Orianos 51 

Othreis 6, 8, 63, 79. so 

pacilicus (Brachyplatys) 5. 50 
pacificus (Ooencyrtus) 5. 13. 49. 50. 1 12 
pallidipcs (AphidencyrtUS, Ooencyrtus) 

5,6,8, 13. 51. 52. 53.62.79. 113. 114 
Papilio 6. 32. 79. 80 
papilionis (Ooencyrtus) 2. 5. 6. 7. 10. 15. 

67, 67, 79, 80. 81. 82. i<}. 130. 131. 132 
papulosa (Tessaratoma) 5. 19 
papuensis (Amblypelta) 4. 82, 83 
Paragus4, 16, 17 
patriciac (Ooencyrtus) 82 
penicillatus (Dolycoris) 5, 78 
pcrpusilla (Pyrilla) 5, 20, 67. 79 
phaeorrhoea (Nygmia) 6, 27 
Phalanta 6. 79, 80 
phalantha (Phalanta) 6. 79. 80 
phoebi (Ooencyrtus) 6, 12. 26. 27 
phongi (Ooencyrtus) 5. 11. 19. 20. 21. 

22.90 
Piezodorus 4. 82, 8 

pilosus (Ooencyrtus) 13, 43. 52, 53, 1 14 
pindarus (Ooencyrtus) 4, 12, 30, 31. 32. 

33, 98, 99 
pinicolus (Ooencyrtus) 2. 46 
piperis (Dasynus) 5. 82, 83 
pityocampae (Ooencyrtus) 2, 42 
plautus (Ooencyrtus) 6. 12. 30, 32. 33. 

100 
Plesispa 4, 31 



Podontia 4. 29, 30 

podontiae (Ooencyrtus. Schcdius) 4, 12, 

29.30.31. 32.98 
polytes (Papilio) 6. 79. 80 
pondcrosum (Pycanum) 5. 19 
populi (Amorpha) 7, 78 
Precis 79. 80 
prolana (Mictis) 5. 82 
promethea (Callosamia) 7. 27 
Pseudaletis 16 
Pseudolitomastix 9 
Pscudotheraptus 5. 8. 82, 83 
Psyllacphagus 9. 16. 17 
pura (Asarkira. Dideopsis) 4. 16, 17 
pulchella (Utethcisa) 5.82 
Pycanum 5. 19 
pyrctorum (Eriogyna) 7, 27 
Pyrilla 5. 20, 67. 79 
pyrillae ( Ageniaspis) 67 
pyrillac (Ooencyrtus) 67 
pyrogramma (Agonoxena) 5. 81 

quatuordecimpunctata 4. 29. 30 

radicornis (Tecoma) 16 
repanda (Taragama) 6. 78 
reichei 4. 31 
Riptortus 4, 62, S2.83 
Rondotia 5. 2') 
rumanzovia (Papilio) 6. 79. 80 

salicis (Stilpnotia) 6, 27 
Schedius 9. 26. 30. 78. 79 
SCUtelll iris (Ischiodon) 4. 16. 17 
segestes (Ooencyrtus) 15. 75. 76. 78. 

127, 128 
selenaria (Ascotis) 8 
sexsignatus (Agrilus) 51 
shadabi (Clavigralla) 4. 83 
shakespearci (Coccidoxenus, 

Ooencyrtus) 5. 16. 80.81 
spectabilis (Dendrolimus) 6. 27 
sphingidarum (Ooencyrtus) 7, 10. 15, 63. 

SO. 81 
spongitus (Anagyropsis) 81 
Stilpnotia 6. 27 
Stauropus 6, 58 

submctallicus (Ooencyrtus) 2. 3. 8. 10 
s\ i phidis (Ooencyrtus) 18 

Tanaccia 6, 80 

Taragama 6. 78 

tartacea (Malacosoma) 6 

Tacoma 17 

telenomicida (Encyrtus, Ooencyrtus) 3. 

5.6,7.8. 15,78'. 79. 130 
Tclenomus 80 
terenellus (Chilo) 5, 79. 80 
Tcrminalia 62 
Tcssaratoma 5. 19 
Tetracnemelhi 9 
Tctrastichus 26 

theobromae (Amblypelta) 8, 79 
thrax (Erionota, Hadart) 6, 8, 51. 52, 79 
Tirumala 5, 79. 80 
tomentosicollis (Clavigralla) 5. 82 
Trichogramma 4 
Trichomasthus 9. 10 
trinidadensis (Ooencyrtus) 3, 8. 83 
tristis (Anasa) 8, 80, 82 



136 D. -W.HUANG AND J.S. NOYES 



Troides 6, 79, 80 4, 5, 8, 16, 70, 79, 80. 82, 83, 132, 133 viridula (Nezara) 5, 8, 80, 82, 83, 84 
Turnaca 6, 65 

valcanus (Ooencyrtus) 14, S3, 54, 115 

unipuncta (Mythimna) 6, 16 vertumnus (Ooencyrtus) 14, 55, 56, 57, 

urania (Ooencyrtus) 14, 54, 55, 115 71, 72, 73, 116 

Utetheisa 5, 82 vesta (Ooencyrtus) 14, 56, 57, 71 , 74, Xesmatia 9, 34 

utetheisae (Aenasioidea, Ooencyrtus) 3, 116 xylina (Lymantria) 6, 27 



wayi (Pseudotheraptus) 5, 8, 82, 83 



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I 



A revision of the Indo-Pacific species of Ooencyrtus (Hymenoptera: 
Encyrtidae), parasitoids of the immature stages of economically 
important insect species (mainly Hemiptera and Lepidoptera) 

D.W. Huang andJ.S. Noyes 



ENTOMOLOGY SERIES 

Vol.63, No. 1, June 1994